On the potential use of magnesium and strontium concentrations as ecological indicators in the calcite skeleton of the red coral (Corallium rubrum)

 The variability of magnesium, strontium and calcium concentrations in the two skeleton types (sclerites and axis) of the red coral (Corallium rubrum) was assessed by using X-ray fluorescence spectroscopy and microprobe analysis as a prerequisite for their use as ecological indicators. Axis cross sections showed light and dark circular growth bands corresponding to fast and slow growth, respectively. Using microprobe analyses the Mg and Sr concentrations ranged from 2.8 to 3.0% and from 0.21 to 0.29%, respectively, in the sclerites and from 2.4 to 3.0% and from 0.1 to 0.28%, respectively, in the axis. Mg/Ca and Sr/Ca ratios varied within sclerites (average, 12% for both ratios) and within the axis of single colonies (average, 20% for Mg/Ca and 48% for Sr/Ca). Sr/Ca ratios in the axis were lower in fast-growing branch tips than in older, more basal parts of the colonies, whereas the Mg/Ca ratio did not differ significantly between colony regions. Mg/Ca and Sr/Ca ratios in the axis decreased significantly with depth, and we estimated an increase of the Mg/Ca ratio of 0.004–0.006 per degree Celsius. In all depth layers, Sr/Ca ratios showed a significant direct relationship with skeleton density in axis cross sections, whereas there was no significant relationship for Mg/Ca. Overall, our data indicate that temperature promotes the incorporation of Mg in C. rubrum as in other calcite skeletons, whereas Sr concentrations are inversely related to growth rate. This preliminary study suggests that Mg and Sr concentrations in the axis of the red coral have a strong potential as ecological indicator for temperature and growth rate. Received: 3 March 2000 / Accepted: 30 August 2000     

Studies on reef corals. III. Fine structural changes of calicoblast cells in Pocillopora damicornis during settling and calcification

The site of reef-coral calcification has been studied in the branching coral Pocillopora damicornis Lamarck. Electron microscopy and X-ray microprobe analysis were performed on the calicoblast epidermis of newly settled larval stages and of adult coral. During settling, the heterogeneous columnar cell composition of the planktonic larva epidermis is replaced by a simple epithelium consisting of a single cell type, the calicoblast cell. Metamorphosis appears tightly linked to settling, with cell changes occurring within hours after attachment, and is marked by the appearance of a new secretory cell. The calicoblast cell of the adult coral is extremely flattened, and interdigitates extensively with adjacent calicoblast cells. This cell possesses a featureless plasma membrane lacking microvilli or flagella. It characteristically contains large membrane-bound vesicles with homogeneously fine granular contents. Preliminary microprobe analysis indicated a higher calcium content in these vesicles than in surrounding tissue; however, not in concentrations suggesting calcium-carbonate precipitation. They may represent sites of organic matrix synthesis. The calicoblast epidermis is separated from the underlying coral skeleton by a narrow gap. This gap appeared devoid of substructure, either organic or inorganic. The coral soft tissues are attached to the skeleton by mesogleal attachment processes, the desmoidal processes. These consist of a complex fibrous network originating in the mesoglea, and inserting onto the skeleton via specialized attachment regions consisting of electron-opaque membranous plaques. Skeletogenesis in reef-corals probably occurs extra-cellularly, external to the calicoblast epidermis, by simple overgrowth of the skeleton.     

Patterns of formation and the ultrastructure of the larval skeleton of Pocillopora damicornis

The growth and development of the tissues and skeleton of settled larvae of the reef coral Pocillopora damicornis (Linnaeus), collected in December 1983 from Ko Phuket, Thailand, were investigated using light microscopy, scanning electron microscopy and transmission electron microscopy. The rate of development of larval skeletons was very variable, preventing the chronological sequencing of skeletal growth. However, four growth stages in the development of a complete larval skeleton from first settlement were identificd: Stage 1, deposition of the first elements of the basal plate upon settlement; Stage 2, completion of basal plate, and deposition of skeletal spines and ridges in positions corresponding to the septal cycles; Stage 3, formation of the corallite wall and septal and costal cycles; Stage 4, the complete larval skeleton which represented the maximum growth attained eight days after settlement. The configuration of the larval tissues, particularly the aboral ectoderm, mirrored the four developmental stages. The deposition of the larval skeleton was correlated with the metamorphosis of the aboral ectoderm from a columnar to a squamous morphology. The basal plate of the larval skeleton had two layers of crystals orientated perpendicular to each other. The architecture of the complete larval skeleton is described and compared to that of the adult skeleton of P. damicornis. The results are discussed with respect to previous concepts of the formation of the larval skeleton of scleractinian corals and coral calcification.     

Structure and function of the locomotory system of the Scyphomedusa Cyanea capillata

The structure and function of the locomotory system of the jellyfish Cyanea capillata L. were studied by dissection, histology, photography and cinematography. The swimming muscle consists of a highly-folded subeptithelial-stratum of striated fibrils arranged on larger mesogleal folds. A proximal coronal muscle divided into 16 fields is anchored to the umbrella proper along the radial septa separating the fields. Sixteen distal couples of radial muscles are anchored to peripheral mesogleal buttresses. On the lower surface of the umbrellar mesoglea is a series of 16 deep radial grooves confluent with a coronal groove. These grooves (joints) are anchored to the exumbrellar surface of the umbrella by rows of large branching mesogleal fibers. During contraction, the muscles cause a folding of the umbrella around the system of joints. Sequences of swimming and turning were analyzed, and it was concluded that this swimming mechanism is much less refined than the hydrozoan mechanism.     

Determination of canthaxanthin in the red coral (<Emphasis Type="Italic">Corallium rubrum</Emphasis>) from Marseille by HPLC combined with UV and MS detection

Corallium rubrum is the most famous and precious coral due to the intense characteristic red colour of its skeleton. We have determined the presence of carotenoids in natural samples of C. rubrum from Marseille, Riou, and investigated their chemical composition. Analysis was performed both on soft tissues and hard tissues including spicules and skeleton. Since hard tissues are made of a mineral fraction and an organic fraction obtained after demineralization, extraction was performed both with and without demineralization by EDTA. The extracts were analyzed by complementary methods of TLC, HPLC/DAD and HPLC/MS. The components were separated by RP-18 chromatography column using acetonitrile/methanol for HPLC/DAD and acetonitrile/water gradient for HPLC/MS analysis. Our results give the first evidence of the presence of canthaxanthin, 4,4′-diketo-β-carotene, as the major carotenoid in all samples. Spicules exhibited higher content in carotenoids than skeleton. Demineralization treatment improved the efficiency of carotenoid extraction by an average factor of 5 and showed that organic matrix contains canthaxanthin.     

Fungi in corals: black bands and density-banding of Porites lutea and P. lobata skeleton

 Dark coloration of coral skeleton forming black bands is commonly observed in fractured, massive-coral colonies (Porites lutea and P. lobata) collected from May- otte Island in the Mozambique Channel and Moorea Island in French Polynesia. Black-banding was similar in corals from the two areas and was associated with an assemblage of microbial endoliths: Ostreobium queketti, a common siphonal chlorophyte, and a type of Aspergillus-like fungus. Fungi of coral skeletons are capable of euendolithic growth entirely within the skeleton, and of cryptoendolithic growth whereby they spread from the skeleton into the skeletal pores. The morphology and size of fungal hyphae differs significantly between euendolithic, cryptoendolithic and reproductive phases. Reproductive phases involve formation of conidiophores. Insoluble residues in black bands involve a dark pigment and a dark membranous veil. When attacked by fungi, the algae are usually destroyed. They darken and are threaded by dense, dark-brown, fibrous excrescences. The fungi excrete a dark pigment that stains the surrounding skeletal carbonate black. The pigment is organic, and its presence correlates with higher concentrations of polysaccharides. Black bands match high-density bands of the coral skeleton. Both black bands and high-density bands form at the end of the rainy season in Mayotte. Thus, black-banding in the corals studied is caused by a series of events, beginning with an increase in the abundance of endolithic algae followed by an increase in skeletal density. The algae are then attacked by fungi, which produce more cryptoendolithic hyphae and conidia that are associated with production of the dark pigment. Received: 29 January 1999 / Accepted: 29 September 1999     

Monthly skeletal extension rates of the hermatypic corals<Emphasis Type="Italic"> Montastraea annularis</Emphasis> and<Emphasis Type="Italic"> Montastraea faveolata</Emphasis>: biological and environmental controls

Monthly skeletal extension rates were measured in colonies of Montastraea annularis and M. faveolata growing at Mahahual and Chinchorro Bank, in the Mexican Caribbean. Temperature, light extinction coefficient (k_d), sedimentation rate, dissolved nutrients and wave energy were used as indicators of environmental conditions for coral growth. Zooxanthella density and mitotic index, nitrogen, phosphorous and protein in coral tissue, and living tissue thickness were measured during periods of high-density-band (HDB) and low-density-band (LDB) formation. To test their value as indirect measures of competition between zooxanthellae and host, as well as coral health and performance in both species, these biological parameters were also measured, during the HDB-formation period, in corals collected at La Blanquilla. This reef is located in the Gulf of Mexico, in an area of suboptimal environmental conditions for coral growth. M. faveolata had a significantly higher skeletal extension rate than M. annularis. Corals growing in Mahahual had significantly higher skeletal extension rate than those living in Chinchorro Bank. This is consistent with inshore–offshore gradients in growth rates observed by other authors in the same and other coral species. This is probably due to less favorable environmental conditions for coral growth in near shore Mahahual, where there is high hydraulic energy and high sedimentation rate. Contrary to observations of other authors, skeletal extension rate did not differ significantly between HDB- and LDB-formation periods for both species of Montastraea. Both species produced their HDB between July and September, when the seawater temperatures are seasonally higher in the Mexican Caribbean. Tissue thickness indicated that environmental conditions are more favorable for coral health and performance during the HDB-formation period. Mitotic index data support the idea that zooxanthellae have competitive advantages for carbon over the host during the LDB-formation period. So, corals, during the LDB-formation period, with less favorable environmental conditions for coral performance and at a disadvantage for carbon with zooxanthellae, add new skeleton with little or no opportunity for thickening the existing one. This results in an equally extended skeleton with lower density, and the stretching response of skeletal growth, proposed for M. annularis growing under harsher environmental conditions, also occurs during the LDB-formation period.Communicated by P.W. Sammarco, Chauvin     

A basidiomycete isolated from the skeleton of<Emphasis Type="Italic"> Pocillopora damicornis</Emphasis> (Scleractinia) selectively stimulates short-term survival of coral skeletogenic cells

Endolithic fungi bore through the extracellular calcium carbonate skeleton of reef-building scleractinian corals, both healthy and dead, and effect net erosion of coral reefs. Potential fungal interactions with coral tissue were investigated using an in vitro approach suggested by earlier observations of skeletal repair cones at the site of fungal perforation in Porites sp. A fungal strain was isolated from the skeleton of a long-term culture of healthy, tissue-covered, Pocillopora damicornis Linnaeus colonies maintained in a recirculating system in Monaco. As coral soft tissue spontaneously dissociated in vitro, the skeleton became exposed and hyaline hyphae emerged radially from 15% of the total clipped branches. In this study, which was performed between January 2001 and March 2003, 35 skeleton–hypha explants were embedded in agar-based solid medium, yielding 60% hyphal growth. A fungal strain (F19-3-1) of the dominant (80%) morphology was isolated and propagated in agar-based solid medium. The strain was identified by 18S and 26S rDNA gene sequence analysis as a basidiomycete in the genus Cryptococcus. Co-cultures were used to provide experimental exposure of coral soft tissue to the fungus. The fungus extended the survival of coral cells by 2 days, selectively maintaining skeletogenic cell types. This effect may be interpreted as stimulation by the fungus of a short-term coral defense response.Communicated by J.P. Grassle, New Brunswick     

Studies on reef corals. I. Skeleton formation by newly settled planula larva of Pocillopora damicornis

Initiation of coral-skeleton formation was studied in the reef-coral Pocillopora damicornis Lamarck. Observations were made on sequential skeletal growth stages of newly settled planula larvae during the first 22 days following settling onto glass microscope slides. Techniques used include phase light microscopy, scanning and transmission electron microscopy, and powder X-ray and selected area electron micro-diffraction. Formation of the skeleton is initiated immediately on settling of the larva. The primary calcareous elements are of two types — flattened spherulitic platelets, and smaller rod-like granules. Rudimentary primary septa are clearly defined within 6 h after settling. Fusion of the primary calcareous elements results in the formation of the larval basal disc within 48 to 72 h. With transmission electron microscopy, this basal disc is found to differ from subsequent adult calcification in (1) considerably lesser degree of mineralization, (2) smaller crystal size, (3) more random orientation of the crystals, and (4) the presence of trace amounts of calcite in addition to aragonite. The basal disc with its septal rudiments constitutes a true larval skeleton, differing in morphology, micro-architecture, and crystal type from the fibrous growth characterizing the adult skeleton.Contribution 419, Hawaii Institute of Marine Biology, University of Hawaii.     

Comparative biometry of Eunicella singularis (Gorgonian) sclerites at East Mediterranean Sea (North Aegean Sea, Greece)

The two forms of Eunicella singularis sclerites, spindles and clubs, were examined in two sites in East Mediterranean Sea, Greece. The comparison of the sclerites size was based on the hydrodynamic profile of the sites (the site of Arethoussa was exposed and the Phidonissi site was sheltered), on the sites bathymetry (AR 5–8 m, AR 9–13 m, PH 5–10 m, PH 11–13 m) and on the position occupied by the sclerites on the gorgonian colonies (top, middle and base). With the usage of image system analysis the following microfeatures were measured or calculated: projection, length, width, elongation factor, compactness factor, length × width and length/width. The present study confirms the hypothesis that spindles are involved in the recovering of the gorgonian colonies and clubs contribute to the flexion capacity of the colonies. The spindles of the shallow E. singularis colonies (AR 5–8 m) were significantly bigger (length, width) compared with the others, in order to assure the best recovery of the colonies after their flexion, induced by higher water current velocity in shallow water. The clubs occupy the intern part of the mesoglea and their size (length, width) increased in colonies from the shallow exposed site in order to facilitate the gorgonian flexion. The present study demonstrates the gorgonian adaptation (in a microscopic scale) to the environmental pressure of hydrodynamic conditions.     

An electron microscope study on the mineralization of the skeleton of the staghorn coral Acropora hebes

The skeleton formation in the reef-building coral Acropora hebes (Dana) was ultrastructurally investigated by observing the skeletogenic tissue and the skeleton in the apical portion of the branches. The skeletogenic tissue was made of a layer of tall calicoblastic cells which displayed high exocytotic activities. A number of hollow spherules and tiny vesicles were found in the sub-epithelial space between the calicoblastic cell layer and the skeletal plate. These organic materials appeared to occur in the perinuclear Golgi vesicles in the cells. The energy-dispersive X-ray analysis revealed a calcium element in the osmiophilic granules of the calicoblastic cells, but not in any other cell organelles. As the granules showed no diffraction pattern, it was suggested that they could be a Ca-reservoir in the cells. Crystalline particles were found to deposit on the periphery of the sub-epithelial spherules. The spherules developed to spherular crystals by depositing granulated crystalline particles. Two of the spherules appeared to fuse with each other to form a spindle-shaped crystal. The spherular and spindle-shaped crystals accumulated on the thecal ridge and the lateral side surface of the thecal plate, and seemed to contribute to the elongation and thickening of the thecal plate. The thecal plate exhibited a porous structure which probably originated from an aggregation of the central cores of these crystals. On the surface of the thecal plate more than about 5 m thick, scale-like structures composed of spherular crystal substructures were observed. These observations suggest that mineralization in A. hebes occurs in the extracellular space by elabolating the spherular and spindle-shaped precursor structures and that growth of the corallite is brought about by an aggregation and coalescence of these crystals.     

Structure,iodination and growth of the axial skeletons of Muricea californica and M. fruticosa (Coelenterata: Gorgonacea)

Axial skeletons of two closely related, sympatric species of gorgonians were examined for details of gross morphology, internal organization and fine structure. Banded fibers with a periodicity of 430 Å and oriented circum-axially to the longitudinal axis of the skeleton (i.e., lying on the plane perpendicular to the axis), were revealed by electron microscopy in the axial skeleton of Muricea californica; they indicate the presence of a collagen-like protein in the gorgoin of this species. No circum-axial fibers were found in the axial skeleton of M. fruticosa. Differences in fine structure are related to gross morphological differences in the skeletons. A sequence of events for the addition of new material to the axial skeleton (growth) is described based on histological observation. Iodine-125 is used as a tracer for the deposition of new skeletal material, and is suggested as a new method for studying gorgonian growth rates.     

Pit structure and trophic relationship of the coral pit crab Cryptochirus coralliodytes

Individuals of the pit crabs Cryptochirus coralliodytes Heller inhabit massive corals of the family Faviidae. Their pit walls were observed to be covered by blue-green algae and fungi. We suggest that the crabs enhance the growth of these algae and fungi with their metabolic excretions, which contain ammonium. The endolithic algae and the fungi may facilitate the abrasion of the coral skeleton by the crabs, by perforating it and thus weakening the skeletal structure. Computerized tomography analysis revealed dense skeletal material around the pits. Transverse sections showed that the calcification around the pit was similar to other parts of the colony, whereas the macro-architecture was different. Such a difference is the result of the crabs' influence on the corals' living tissue, possibly on the calicoblast which deposits the coenosteum. Crabs, which were exposed to carbon-labeled corals for 1, 7 and 18 d, accumulated labeled carbon, indicating transfer of carbon from the coral tissue to the crabs. Histochemical examination of the stomach and gut of crabs revealed the presence of mucopolysaccharids in the gut, supporting the hypothesis that the crabs eat coral products. The findings of this study provide additional evidence that C. coralliodytes are parasites and support the general hypothesis that a nutritional relationship may have served as a basis for selection. Received: 20 October 1998 / Accepted: 29 April 1999     

Embryogenesis and acquisition of algal symbionts by planulae of Xenia umbellata (Octocorallia: Alcyonacea)

Early embryogenesis of the internally brooding soft coral Xenia umbellata and acquisition of algal symbionts in the course of its planular ontogenesis have been examined by scanning and transmission electron microscopy and by light microscopy. The endoderm of adult X. umbellata harbours symbionts mainly in the tentacles and in the peripheral solenia system. The colonies are gonochoric brooders. Algal symbionts were never found in the sperm sacs, and were only rarely found in the follicular tissue enclosing the oocytes. Fertilized eggs pass into endodermal brood pouches where embryogenesis occurs. Cleavage is holoblastic and leads to formation of a solid blastula. Algal symbionts are conspicuously embedded in the parental mesoglea that coats the young embryo, most probably transmitted by surface adherence. At a further stage, this integument disappears and the algae reside extracellularly among the cells of the newly-formed blastula. After subsequent cell proliferation developing planulae possess an inner mass of yolk-laden cells that contain numerous symbiotic algae. Gradually the yolk disintegrates, leaving a cavity enclosed by ectoderm, a thin mesoglea and an inner endoderm with intracellular symbionts. The mature planulae have already been provided with numerous intracellular symbionts by the time they are expelled from the brood pouches. The markedly early symbiont acquisition by the embryos of X. umbellata may help support their developmental requirements in the course of planular ontogenesis.     

Reproduction in Anthelia glauca (Octocorallia: Xeniidae). II. Transmission of algal symbionts during planular brooding

The soft coral Anthelia glauca Lamarck, 1816, of the family Xeniidae, is found on the reefs of KwaZulu-Natal, South Africa. Its gastrodermal cells contain numerous endosymbiotic unicellular algae (zooxanthellae). A. glauca is a gonochoric species that simultaneously broods its planulae within the pharyngeal cavity of the polyps. Symbiotic algae appear with zygote formation within the pharynx, embedded in amorphous material. The algal cells adhere to the ciliated ectodermal surface of immature planulae and are most probably endocytosed by them. Zooxanthellae are translocated towards the basal part of the ectoderm. Gaps are subsequently opened in the mesoglea into which symbionts surrounded by ectodermally derived material, including plasma membrane, pass. The basal membrane of endodermal cells disintegrates, and the algae bulge into spaces formed in the underlying endoderm. Throughout the process, each zooxanthella resides within a vacuolar membrane in the detached ectodermal cytoplasm. The acquisition process is essentially one in which zooxanthellae are translocated from the pharyngeal cavity into the ectoderm and then through the mesoglea into the endoderm, culminating in the final symbiotic state. The direct transmission of symbiotic algae to the eggs or larvae probably provides the most efficient means whereby zooxanthellae are acquired by the host progeny. Received: 15 July 1997 / Accepted: 25 February 1998     

Short-term growth measurements of corals using an accurate buoyant weighing technique

An accurate method for determining the growth rates of the skeleton of isolated branch tips (nubbins) of corals over intervals of less than 24 h is described. The skeletal weight of the coral was estimated from its buoyant weight in seawater whose density had been accurately determined. The coral tissues accounted for between 1 and 5% of the total buoyant weight in Pocillopora verrucosa and Acropora humilis with differing relative tissue biomass. After correcting for tissue buoyant weight, predictions of skeletal weight were accurate to within 1%. The method was used to estimate the growth of sample nubbins of Porites porites of similar diameter, in 2 m of water at Discovery Bay, Jamaica. Since growth of these branch tips is apical, growth rate could be expressed without correction for the size. The mean 24 h skeletal growth rate ranged between 40 and 47 mg. Differences could be measured between day-time and night-time growth, the day: night ratio being 3.7. The method also showed that P. porites virtually ceases calcification during the 4 to 5 d periods that it becomes enclosed in a mucus tunic. Nubbins of P. porites attached to the reef at different locations showed clear differences in growth rate with depth, and between clear and turbid water sites. The growth rate of nubbins was compared with that of branch tips of whole corals by measuring the linear extension after staining with Alizarin Red S. After 3 1/2 mo, the mean linear extension was 4.1 mm in each case, indicating that the growth rate of nubbins is the same as that of branch tips of the whole colony. It is suggested that this buoyant weighing technique will find applications in laboratory experiments with calcification mechanisms and as a bioassay on reefs exposed to environmental stress.Contribution No. 464 of the Discovery Bay Marine Laboratory     

Population structure of an exploited benthic cnidarian: the case study of red coral (Corallium rubrum L.)

Octocorals are an important part of many ecosystems as they add three-dimensional complexity to the benthos and thereby increase biodiversity. The Mediterranean red coral (Corallium rubrum, L. 1758) is a longevous octocoral that is harvested commercially, yet natural and anthropogenic influences on its population size structure are little understood. This study found that some harvested red coral populations had a significantly different size structure when compared to populations at the nearby Marine Protected Area (MPA) of Medas Islands at the Spanish Costa Brava (NW Mediterranean). Eighty-nine percent of the red corals in the harvested Costa Brava area are less than 10 years old and 96% of all colonies have not yet grown more than second-order branches. The size/age distribution of the harvested population is notably skewed towards younger and smaller colonies. Thus, although red coral is still abundant, its population structure is strongly distorted by harvesting. The results confirm that MPAs are useful to distinguish between anthropogenic and natural influences on population structure. However, 14 years of protection appears to be an insufficient recovery time for a longevous octocoral population such as red coral.     

Photobiology of endolithic microorganisms in living coral skeletons: 1. Pigmentation, spectral reflectance and variable chlorophyll fluorescence analysis of endoliths in the massive corals Cyphastrea serailia, Porites lutea and Goniastrea australensis

We used microscopy, reflectance spectroscopy, pigment analysis, and photosynthesis-irradiance curves measured with variable fluorescence techniques to characterise the endolithic communities of phototrophic microorganisms in the skeleton of three massive corals from a shallow reef flat. Microscopic observations and reflectance spectra showed the presence of up to four distinct bands of photosynthetic microorganisms at different depths within the coral skeleton. Endolithic communities closer to the coral surface exhibited higher photosynthetic electron transport rates and a green zone dominated by Ostreobium quekettii nearest the surface had the greatest chlorophyll pigment concentration. However, Ostreobium was also present and photosynthetically active in the colourless band between the coral tissue and the green band. The spectral properties and pigment density of the endolithic bands were also found to closely correlate to photosynthetic rates as assessed by fluorometry. All endolithic communities were extremely shade-adapted, and photosynthesis was saturated at irradiances <7 μmol photons m⁻²s⁻¹.     

Variability of stable isotopes and maximum linear extension in reef-coral skeletons at Kaneohe Bay, Hawaii

Stable-isotope and growth records of coral skeletons are often used to reconstruct tropical paleoclimate, yet few surveys have systematically examined the natural variability in coral skeletal ¹³C, ¹⁸O and maximum linear skeletal extension (MLSE) across depth. Here, interspecific, intraspecific, and geographical variations in coral skeletal ¹³C, ¹⁸O, and MLSE were examined in the corals Porites compressa, P. lobata, and Montipora verrucosa grown at 1.7, 5.0, and 8.3 m depth from August 1996 to March 1997 at The Point Reef and Patch Reef #41 field sites in Kaneohe Bay, Hawaii. Coral skeletal ¹³C values significantly decreased with depth and differed between species, but did not vary between field sites. ¹⁸O values were not significantly different across depth within a species, but did differ among species and field sites. High-resolution analysis of the intra-annual variation in skeletal ¹³C and ¹⁸O in P. compressa at 2.0 m depth confirms that these isotopes reflect changes in solar irradiance and temperature, respectively. Changes in MLSE across depth were consistent within, but highly variable among, species. Peak MLSE occurred at 1.7, 5.0, and 8.3 m for P. lobata, P. compressa, and M. verrucosa, respectively. Such interspecific variation in MLSE patterns may be attributable to one or more of the following: increases in zooplankton in the diet, changes in metabolic processes, or changes in growth form with depth. Overall, these results imply that natural inter- and intraspecific variability in coral skeletal ¹³C, ¹⁸O, and MLSE should be considered when interpreting and comparing coral-based tropical paleoclimate data from various coral species, depths, and field sites. Received: 6 October 1998 / Accepted: 8 July 1999     

Associations between metals and the blue mesogleal protein of Cassiopea xamachana

The blue mesogleal pigment of the symbiotic jellyfish, Cassiopea xamachana Bigelow, 1882, is composed of two subunits, a larger glycosylated (35 kDa) moiety and a non-glycosylated (30 kDa) variant in lower concentration. In solution, the subunits assemble in large complexes of at least 10⁶ kDa. The pigment, known as Cassio Blue, appears to mitigate excessive solar radiation while allowing the passage of the wavelengths optimal for photosynthesis by the numerous algal symbionts in the mesoglea of the jellyfish. The pigment is an abundant protein comprising about 6% of all animal protein in the whole jellyfish and about 33% of all animal protein in the oral appendages. The protein also contains a diverse array of metals, notably Ag, Ca, Cu, Fe, Mg, and Zn, with traces of others. Metal stoichiometry varies among isolates averaging about 1 mol of all metals, taken together, for each mole of the pigment. Given the broad array of metals present, the pigment may also serve another purpose, for example, as a metal reservoir or trap. Few other proteins are associated with such a spectrum of metals. In addition, the amino acid sequences of the pigment tryptic peptides have no reasonable matches in any of the sequence databases. Our findings, taken as a whole, suggest that the Cassio pigment is indeed unusual and is likely a representative of a novel category of proteins, the original member of which is rpulFKz1, a chromoprotein endowed with Frizzled and Kringle domains.