Functional autotrophy of Heteroxenia fuscescens (Anthozoa: Alcyonaria): carbon assimilation and translocation of photosynthates from symbionts to host

The association of the alcyonarian Heteroxenia fuscescens (Ehrb.) with its cytosymbiotic algae shows structural and physiological adaptations optimizing the living together of the two partners as one functional unit. To enhance the energetic contribution of the autotrophic partner, the organization of the heterotrophic partner bears typical plant-like imprints. Up to 20% of the inorganic C photosynthetically fixed was translocated to the host (=2 mg C mg d.w.^-1 d^-1). This net C gain by the host is used for anabolic purposes including the deposition of storage material. Especially the wax-esters and triglycerides of the host-in contrast to those of the symbionts-were intensively labelled. The in-vivo ¹⁴C-fixation of zooxanthellae is more than double the in-vitro fixation. In both symbionts and host, the lipids show the highest relative ¹⁴C-incorporation. In particular the polyol component was strongly labelled. After 120 min of continuous incubation, approximately 40 labelled intracellular metabolites were detectable in the ethanol/water soluble fractions of zooxanthellae. Glycerol is the main low-molecular weight carbohydrate being transferred. This is corroborated by the deacylation of lipids of the host.     

Requirement of exogenous inducers for metamorphosis of axenic larvae and buds of Cassiopea andromeda (Cnidaria: Scyphozoa)

Planula larvae and asexually-produced buds of the rhizostome scyphozoan Cassiopea andromeda (collected throughout the year in Eilat, Israel) have the ability, under axenic conditions, to attach to a substrate and undergo morphogenetic development to form a polyp (=scyphistoma) in: (1) the presence of unidentified inducers found in the adult habitat and (2) the presence of cefined organic compounds. Axenic planulae and buds were unable to settle and complete metamorphosis in autoclaved artificial or natural seawater from the North Sea when maintained without food, but continued swimming while decreasing in size and protein content, eventually dying within three months. When maintained in autoclaved seawater from the Red Sea, between 25 and 46% of the planulae and 4 and 11% of the buds metamorphosed within 30 d. Axenic solutions of cholera toxin, thyroid stimulating hormone, and pancreatic casein hydrolysate peptides in artificial seawater induced morphogenic development of 20 to 100% of planulae and buds within 2 to 18 d. The natural inducer(s) in Red Sea seawater, though unidentified, may have characteristics similar to the large proteins and small peptide inducers used in this study. Planulae and buds older than 20 d metamorphosed sooner and responded to lower concentrations of pancreatic casein hydrolysate peptides than younger individuals. This may be a physiological mechanism for enhancing metamorphosis and survival in nature. The data show that settlement and metamorphosis can be induced by solutions of cholera toxin and thyroid stimulating hormone, suggesting that, as in mammalian systems, the mechanism of action of these chemicals may involve cyclic adenosine monophosphate (cAMP) as an intermediate messenger. However, dibutyric cAMP, which is capable of passing through membranes and functioning normally inside the cell, did not induce metamorphosis of buds, and the levels of intracellular cAMP in buds and larvae typically increased slowly during induction of metamorphosis, unlike the high and rapid increases associated with cAMP-mediated biochemical events in mammalian cells. These results suggest that the observed cAMP changes seen were associated with metamorphic development, but not with the triggering mechanism.     

Development and survivorship of zooxanthellate and azooxanthellate primary polyps of the soft coral Heteroxenia fuscescens: laboratory and field comparisons

Marine Biology - The zooxanthellate Red Sea soft coral Heteroxenia fuscescens releases aposymbiotic planulae nearly all year round, with higher numbers in the summer than in other seasons. After...     

Reproduction and development of the blue coralHeliopora coerulea (Alcyonaria: Coenothecalia)

Up to the present time, the reproductive cycle and larval development ofHeliopora coerulea, living fossil and sole member of the alcyonarian order Coenothecalia, have been undescribed. Populations ofH. coerulea studied on the Great Barrier Reef between 1984 and 1989 were found to have separate sexes and an annual gametogenic cycle. Mature eggs were large (>800µm diam), and their development was initiated internally. There appeared to be a lunar or semi-lunar periodicity in reproductive behaviour, with brooding of larvae commencing around either full or new moon. Development proceeded via superficial cleavage and a solid stereoblastula. Subsequent development took place on the surface of the colony, within a temporary brood-space created by protracted distension of the polyps. Larvae lacked zooxanthellae and were brooded for between 6 and 14 d prior to leaving the colony. Reproduction and development reflected the taxonomic affinities ofH. coerulea, but brooding and embryological development also appeared to be constrained by its morphology. Reproductive and larval characteristics ofH. coerulea may offer a partial explanation of the current biogeographic range ofH. coerulea.     

Bioluminescence of deep-sea coronate medusae (Cnidaria: Scyphozoa)

Bioluminescence is the production of visible light by a living organism. The light commonly appears as flashes from point sources (involving one or more cells, usually described as photocytes) or as a glandular secretion. A visible flash usually involves synchronous light emission from a group of cells or, if from a single-celled organism such as a dinoflagellate, from a group of organelles. The number of cells (or organelles) responding synchronously is the main determinant of the flash intensity. Bioluminescence is a common phenomenon in many deep-sea animals and is widespread among the Cnidaria. In this paper, we compare and contrast in situ and laboratory recordings of the bioluminescent responses of specimens of the deep-sea scyphozoans Atolla wyvillei, Atolla vanhoffeni, Atolla parva, Nausithoe rubra, Paraphyllina intermedia, Periphyllopsis braueri and Periphylla periphylla. Displays in all seven species consist of localised flashes and propagated waves of light in the surface epithelium. The first few single waves propagate at rates of up to 60 cm s^-1 but subsequent ones in any sequence of stimuli gradually decrease in speed. After several single wave responses, a subsequent stimulus may elicit multiple waves that persist for several seconds. Following such a frenzy, the specimen becomes temporarily refractory to further stimuli, but if rested will recover its normal responses and may produce further frenzies. The dome area, situated above the coronal groove, of the genera Paraphyllina, Periphylla, and Nausithoe is covered with luminescent point sources. Such point sources are generally absent from the dome of species of Atolla. Captured specimens of A. parva also produce secretory bioluminescence, corroborating prior in situ observations of this ability. Secretory bioluminescence in P. periphylla takes the form of scintillating particles released from the lappet margins. We did not observe secretory displays in specimens of any other species in the laboratory, but one instance of apparent secretory luminescence was recorded in situ in a specimen of A. wyvillei.Communicated by J. P. Thorpe, Port Erin     

Population dynamics of Eudendrium glomeratum (Cnidaria: Anthomedusae) on the Portofino Promontory (Ligurian Sea)

Eudendrium glomeratum Picard, in the Ligurian Sea, is one of the major components of hard-bottom sessile zoobenthos in the cold season. It settles mainly between 10 and 40 m depth, forming a seasonal facies. The presence of E. glomeratum has been evaluated by measuring in situ the height of the colonies present within a standard surface of 1 m². Observations were carried out every other week for a whole year. The height of the colonies was correlated with their biomass (wet weight) and trophic functionality (number of polyps). The quantitative evaluation of the population shows that the situation is stable, with almost identical values at the extremes of the observed yearly time series. The species is present from October to April–May, and it is absent in the summer. The formation and disappearance of the E. glomeratum population are sudden. The colonies are sexually mature from their appearance in October to December–January. Different categories of colonies are recognizable; and are essentially referable to two groups: (1) colonies deriving from regeneration of colonies present in the previous year (45%); (2) colonies deriving from planula settlement (43%). A third group (12%) is of uncertain origin and may belong to either of the abovementioned categories. Mortality is 60% in the first group, and 82% in the second one. By dead colonies is meant those which do not reappear after a yearly cycle.     

The morphogenesis of ephyra in Coronatae (Cnidaria,Scyphozoa)

The morphogenesis of ephyrae of Atorella vanhoeffeni Bigelow, 1909 (Werner, 1967) and of Nausithoe maculata Jarms, 1990 during strobilation is described. We found differences in the developmental pattern of marginal structures and structural changes of longitudinal muscle tubes in particular. During strobilation the polyp’s tetraradiate symmetry is passed to the ephyra of both taxa as a tetraradiate central symmetry that we consider a major plesiomorphic character. The present results also indicate divergent patterns of ephyra development during strobilation that lead to variable marginal symmetries of ephyrae and thus of medusae. Therefore, the marginal symmetry of medusae of N. maculata is octoradiate and of A. vanhoeffeni is hexaradiate. We conclude the latter is stated as derived pattern. These results lead us to maintain both families Nausithoidae and Atorellidae.     

Sexual reproduction in the coral Pocillopora verrucosa (Cnidaria: Scleractinia) in KwaZulu-Natal, South Africa

The mode of reproduction in Pocillopora verrucosa Ellis and Solander, 1786 varies between geographically isolated regions. This scleractinian coral is common along the KwaZulu-Natal coast, and its reproductive mode and period of reproduction were assessed using histological preparations. The study was undertaken from 1992 to 1994 and showed that P. verrucosa is a simultaneous hermaphrodite and broadcast spawner in KwaZulu-Natal. Gametogenesis occurs from October to January, with the gametes maturing simultaneously so that spawning can be synchronised at new moon in January (mid-summer). Zooxanthellae are present in the mature oocytes. Received: 10 September 1997 / Accepted: 6 July 1998     

Reproduction of Dendronephthya hemprichi (Cnidaria: Octocorallia): year-round spawning in an azooxanthellate soft coral

Sexual reproduction in the azooxanthellate octocoral Dendronephthya hemprichi Klunzinger, 1877 was studied in Eilat (Red Sea) for 2 years beginning March 1989. D. hemprichi is a gonochoric broadcasting species. Gonads at all developmental stages were found throughout the year. Small-sized oocytes and sperm sacs, 51 to 100 μm in diameter, are highly abundant, accompanied by numerous smaller primordial gonads. These features result from continuous gametogenesis, and lead to year-round oocyte and sperm maturation and release. Gamete release was observed in the laboratory on all monitoring dates, and most of the female colonies spawned repeatedly for several successive nights. The reproductive features of D. hemprichi constitute an exception to the generalization that free-spawning reef-corals have brief and synchronized broadcast-spawning episodes. This species demonstrates early onset of first reproduction, which further increases its reproductive output. Received: 7 May 1997 / Accepted: 15 May 1997     

Carbon metabolism and strobilation in Cassiopea andromedea (Cnidaria: Scyphozoa): Significance of endosymbiotic dinoflagellates

Scyphopolyps and scyphomedusae of Cassiopea andromeda Forskål (Cnidaria, Scyphozoa) containing dinoflagellate endosymbionts (zooxanthellae) were investigated for rates and pathways of carbon fixation. Photosynthesis by the algae, accounting for 80 and 15 mol C h^-1 on a dry weight basis in medusae and polyps, respectively, by far exceeds dark incorporation of inorganic carbon by the intact association. Photosynthetic carbon fixation is operated via C₃ pathway of carbon reduction. DCMU-treatment (1×10^-6 M and 1×10^-5 M) completely inhibits light-dependent carbon assimilation. Major photosynthates presumably involved in a metabolite flow from algal symbionts to animal tissue are glycerol and glucose. A total of 5–10% net algal photosynthate appears to be seleased in vivo to the host. This is probably less than the energy supply ultimately required for the nutrition of the polyps and medusae. The presence of zooxanthellae proved to be indispensable for strobilation in the scyphopolyps. However, photosynthesis by algal symbionts as well as photosynthate release is obviously not essential for the initiation of ephyrae as is shown by DCMU-treatment, culture in continous darkness, and aposymbiotic controls. It is therefore concluded that strobilation is supported, but not triggered by algal photosynthetic activity. The induction of strobilation thus seems to depend on a more complex system of regulation.     

Use of electrical stimulation for discharging cnidom components of a species of Cerianthus (Anthozoa: Cnidaria)

Direct-current electrical stimulation was used as a means of discharging cnidom components from the marginal tentacles of a species of Cerianthus from Southern Queensland, Australia, in order to facilitate the study of cnidom components by light microscopy. The cnidom consists of b-rhabdoids (3 types), spirocysts, and ptychocysts. Threshold pulses of between 1 and 2 V with a frequency of 1 pulse per second for 2 sec discharge all types of cnidae. However, large numbers of certain cnida types discharge after a longer period of stimulation.     

Strobilation,budding and initiation of scyphistoma morphogenesis in the rhizostome Cassiopea andromeda (Cnidaria: Scyphozoa)

Scyphistomae of Cassiopea andromeda Forskål, 1775 containing symbiontic zooxanthellae did not develop medusae at a constant temperature of 20°C, but monodisc strobilation was initiated after transfer of the polyps to 24°C. After release of the ephyrae and regeneration of the hypostome and tentacular region, the recovered polyps either produced vegetative buds or entered a new strobilation phase. Formation of motile, planula-like buds was not found to be indicative of unfavourable environmental conditions. Intensity of budding was positively correlated with available food and with increase of temperature. Budding was negatively correlated with the number of polyps maintained per dish and with the conditioning of the sea water. Under optimal feeding and temperature conditions, polyps could simultaneously produce chains of buds at 2 to 4 budding regions. Settlement and development of buds into scyphistomae was suppressed in pasteurized sea water and in pasteurized sea water containing antibiotics, but polyps developed from buds in the presence of algal material taken from the aquarium, debris or egg shells of Artemia salina, or on glass slides which had been incubated in used A. salina culture medium. Several species of marine bacteria were detected after staining these slides. One, a Gramnegative coccoid rod, which was identified as a nonpathogenic Vibrio species, was isolated, cultivated as a pure strain, and was proved to induce the development of C. andromeda buds into polyps. Millipore filter-plates coated with Vibrio sp. cells grown in suspension culture were ineffectual, but diluted filtrate initiated polyp morphogenesis. The inducing factor is obviously not a constituent of the bacterial cell surface, but is a product of growing Vibrio sp. cells released into the medium. This product was found to be relatively heat-stable and dialyzable. As to the basic mechanism involved in the induction of polyp formation, it is suggested that the inducing factor (s) acts bimodally by inducing pedal disc development and by eliminating a head inhibitor originating from the basal end of the bud. The life history, and various aspects of medusa-formation and of vegetative reproduction in scyphozoans are reviewed and discussed with particular reference to rhizostome species. Special attention has been paid to some reports of larval metamorphosis controlled by marine bacteria.     

Isolation and cloning of DNA from somatic tissue of soft corals (Cnidaria: Octocorallia)

A method is described for isolating high molecular weight DNA from somatic tissue of soft corals. The method avoids problems associated with the presence of nucleases, pigments and other secondary metabolites in soft corals. Tissue is frozen in liquid nitrogen, pulverized with a pestle and mortar, and the powder extracted with buffered sodium dodecyl sulphate at 4°C in the presence of phenol/chloroform. This technique has been applied toAlcyonium, Sinularia, Sarcophyton andLobophytum species. Repeated sequences have been cloned fromAlcyonium sp., and used to probe slot-blots of genomic DNA and Southern blots of restriction digests. Homologous repeated sequences were detected in threeAlcyonium sp., but not in threeSinularia sp., despite these genera being closely related.     

Population dynamics of Eudendrium racemosum (Cnidaria, Hydrozoa) from the North Adriatic Sea

Benthic suspension feeders in shallow waters develop in relation to the food availability and the variation of physical parameters giving rise to complex communities that act as a control factor on the plankton biomass. The aim of the work is to establish the role of the hydrozoan Eudendrium racemosum in the energy transfer from the plankton to the benthos in marine food chains of the North Adriatic Sea. This study highlighted that the hydroid biomass changed over time in relation to temperature and irradiance, and the highest abundance was observed during summer with about 400,000 polyps m⁻² (about 19 g C m⁻²). The population suffered an evident summer decrease in relation to a peak of abundance of its predator, the nudibranch Cratena peregrina, whose adult specimens were able to eat up to 500 polyps day⁻¹ and reached an abundance of 10 individuals m⁻². The gut content analysis revealed that the hydroid diet was based on larvae of other benthic animals, especially bivalves and that the amount of ingested preys changed during the year with a peak in summer when it was estimated an average predation rate of 13.7 mg C m⁻² day⁻¹. In July, bivalves represented over 60 % of the captured items and about 18 mg C m⁻² day⁻¹. Values of biomass of E. racemosum are the highest ever recorded in the Mediterranean Sea, probably supported by the eutrophic conditions of the North Adriatic Sea. Moreover, our data suggest that settling bivalves provide the greater part of the energetic demand of E. racemosum.     

Effect of nutrient enrichment on the complementary (secondary) metabolite composition of the soft coral Sarcophyton ehrenbergi (Cnidaria: Octocorallia: Alcyonaceae) of the Great Barrier Reef

 A long-term study of the effects of nutrient enrichment on coral reefs (ENCORE Experiment) was carried out at One Tree Island (23°30′S; 152°96′E), Great Barrier Reef, between 1992 and 1996. The experiment involved the addition of water-soluble nutrients to 12 microatolls which contained a range of organisms and were situated within the larger lagoon. Three replicates of each of three nutrient treatments (nitrogen, phosphorus, nitrogen plus phosphorus) and an untreated set of three control atolls were prepared using the 12 selected microatolls. As part of the larger ENCORE experiment, changes in the chemical composition of the alcyonacean soft coral Sarcophyton ehrenbergi Marenzeller (Octocorallia: Alcyonacea), placed in the treatment microatolls, were monitored for a 1 yr period in an attempt to detect any responses attributable to nutrient enrichment. Multivariate analyses were performed to determine whether there were any patterns of response in the different nutrient treatments. At the level of individual metabolites, there were no clear treatment effects. However, the ratio of bioactive or stress metabolites (terpenes) to energy storage metabolites (lipids), referred to as the “physiological-change indicator”, revealed effects of nutrient enrichment. Nitrogen enrichment resulted in a trend towards higher physiological-change indicators than control or phosphorus treatments in the majority of cases, while phosphorus enrichment significantly decreased the ratio relative to controls. In most cases, the physiological-change indicator increased in soft soft corals relocated into contact with the scleractinian Pocillopora damicornis. The potential of soft corals to serve as indicators of a changed nutrient regime is discussed. Received: 23 November 1998 / Accepted: 23 September 1999     

Foraging, prey capture, and gut contents of the mesopelagic narcomedusa Solmissus spp. (Cnidaria: Hydrozoa)

Narcomedusae are the most common group of medusae in the mesopelagic depths of Monterey Bay, California. Numerous capture events of various prey taxa were recorded in situ and analyzed using the Monterey Bay Aquarium Research Institute's remotely operated vehicle "Ventana". In situ video analysis of the stomach contents of 82 Solmissus incisa and S. marshalli revealed 88 identifiable prey items. Most (88%) were gelatinous animals. Of these gelatinous prey, 60.3% were ctenophores, 20.5% were cnidarians, 12.8% were salps, 3.8% were chaetognaths, and 2.6% were polychaetes. Euphausiids accounted for 11.4% of the diet, but they were probably captured adventitiously, as the gut contents of ctenophore prey. The tentacle-first foraging behavior of the narcomedusae is an effective way to capture large, comparatively fast-moving prey, because the fluid disturbance caused by swimming is decoupled from the area of prey capture. This behavior contrasts with the prevailing models of feeding behavior in medusae. Stealth predation may be the dominant mode of capturing prey by medusae in the mesopelagic depths of the oceans.     

Chemical aspects of mass spawning in corals. II. (-)-Epi-thunbergol,the sperm attractant in the eggs of the soft coral Lobophytum crassum (Cnidaria: Octocorallia)

Sea turtles migrate between feeding and nesting areas that are often geographically separated by hundreds or thousands of kilometers. Observations of their aggregations at sea and at nesting beaches have led to the hypothesis that sea turtles migrate in socially structured groups. While this migratory strategy is common to many marine vertebrates, socially facilitated behavior is not well documented in testudines. In 1990 and 1991, we attached satellite transmitters to olive ridleys (Lepidochelys olivacea Eschscholtz) found ovipositing together during a mass nesting at Nancite Beach, Costa Rica, to determine whether they migrate independently or in groups after they leave the nesting beach. Results showed that the turtles were not spatially associated during the internesting period, were capable of re-establishing themselves as a group during a subsequent nesting emergence, and were not spatially associated during their postnesting migrations to oceanic feeding areas. We suggested that what appear to be socially structured groups of L. olivacea are in fact individual turtles simultaneously using the same habitat.     

Effects of temperature on success of (self and non-self) fertilization and embryogenesis in Diploria strigosa (Cnidaria, Scleractinia)

The effects of seawater temperature on the degree of success of self-fertilization versus cross-fertilization and on embryogenesis were investigated in the scleractinian coral Diploria strigosa (Dana 1846). Gametes from nine colonies were collected from the Flower Garden Banks reefs, a set of coral reefs in the northern Gulf of Mexico, ~110 nm offshore from Texas, United States. Gametes from the coral colonies were combined in all possible combinations and placed in 30°, 31°, and 32°C water baths on successive nights, respectively. Each cross, whether selfed or out-crossed, and each temperature treatment, was replicated twice. A high frequency of successful self- and cross-fertilization was observed at all temperature treatments and in both blocks. Higher temperatures, however, commonly produced numerous developmental aberrations during embryogenesis of the larvae. Thus, although fertilization rates can remain high under high temperature conditions, if temperatures remain high for several days, embryonic development and larval viability may be expected to decrease dramatically. We propose that the success of coral larval development may be diminished in areas where abnormally high sea surface temperatures occur during the spawning season. We also propose that highly successful selfing may enhance the abundance of locally adapted genotypes, which in turn may be advantageous where reefs, such as the Flower Garden Banks, are relatively geographically isolated.