Chemical aspects of mass spawning in corals. I. Sperm-attractant molecules in the eggs of the scleractinian coral Montipora digitata

This paper provides the first evidence for sperm chemotaxis in the Scleractinia. Montipora digitata Dana, 1845 (Scleractinia: Coelenterata) is a hermaphroditic coral which reproduces bi-annually, releasing egg-sperm bundles during the mass spawning at Magnetic Island (19°10S; 146°52E) in late spring-early summer, and autumn each year. The buoyant egg-sperm bundles float to the surface where they break apart, releasing eggs and sperm into the ocean. Fertilisation occurs after 30 min. Unfertilized eggs were collected, washed free of sperm, and freeze-dried. The eggs were extracted with dichloromethane, fractionated by chromatography on silica gel, and the fractions assayed for their ability to attract M. digitata sperm. The active fraction was further fractionated by high-performance liquid chromatography, resulting in the isolation of three highly unsaturated fatty alcohols: (1) dodeca-2,4-diynol; (2) tetradec-13-ene-2,4-diynol; (3) (14Z)-heptadeca-14,16-diene-2,4-diynol. Of these three compounds, only Compound 1 attracted sperm of M. digitata. Synthetic Compound 1, produced from simple precursors by known reactions, possessed sperm-attracting activity comparable to the naturally derived attractant. Preliminary experiments suggest that the natural mixture of Compounds 1, 2 and 3 in the ratio 1:4:9 is more effective in attracting sperm from M. digitata than sperm from other Montipora species. Sperm attractants may act to reduce the incidence of hybridisation between different species of Montipora.     

Effects of diterpenes derived from the soft coral Sinularia flexibilis on the eggs,sperm and embryos of the scleractinian corals Montipora digitata and Acropora tenuis

Sinularia flexibilis Quoy and Gaimard, 1833 is a dominant soft coral on many Indo-Pacific coral reefs, and has been found to release toxic compounds (diterpenes), which cause tissue necrosis and death in nearby scleractinian corals. This study investigates how S. flexibilis-derived diterpenes inhibit the development of Acropora tenuis Dana, 1846 and Montipora digitata Dana, 1846 eggs and larvae in vitro. Collection and experimental sites at Magnetic (146°49'E; 19°8'S) and Orpheus (146°28'E; 18°32'S) Islands, Queensland, Australia, were utilized during the spawning seasons of 1989–1992. Freshly spawned coral eggs were placed in solutions of three different terpenes, flexibilide, dihydroflexibilide and sinulariolide, at 5 and 10 ppm, before, during, and after fertilisation. The majority of eggs which were fertilised in the presence of the diterpenes lost their cellular integrity and burst just a few hours after treatment. Terpenes were not toxic to unfertilised eggs, nor to 24 h-old embryos, although sperm ceased swimming activity after 1 h of treatment. The terpenes were not fatal to the sperm because fertilisation still occurred in their presence. The ability of the soft coral-derived diterpenes to inhibit cell division suggests that they may have potential applications in cancer chemotherapy.     

Synchronous spawnings of 105 scleractinian coral species on the Great Barrier Reef

Following observations of mass spawning of hermatypic corals on the Great Barrier Reef in 1981 and 1982, spawning dates were successfully predicted and documented at five reefs on the Central and Northern Great Barrier Reef in 1983. During the predicted times, 105 species from 36 genera and 11 families were observed to spawn. Of these, 15 species were shown to have an annual gametogenic cycle. All but two of the species observed during mass spawnings shed gametes which underwent external fertilization and development. Synchronous spawning was observed both within and between the five reefs studied, which were separated by as much as 5° of latitude (500 km) or almost a quarter of the length of the Great Barrier Reef. The mass spawning of corals took place on only a few nights of the year, between the full and lastquarter moon in late spring. Maturation of gametes coincided with rapidly rising spring sea temperatures. Lunar and diel cycles may provide cues for the synchronization of gamete release in these species. The hour and night on which the greatest number of species and individuals spawned coincided with low-amplitude tides. Multispecific synchronous spawning, or mass spawning, of scleractinian and some alcyonacean corals represents a phenomenon which is, so far, unique in both marine and terrestrial communities.     

Genetic structure of two species of Montipora on a patch reef: conflicting results from electrophoresis and histocompatibility

Spatial patterns of graft acceptance and rejection were different for two species of the coral Montipora from Kaneohe Bay, Hawaii. Electrophoretically distinct tissues were capable of fusing, indicating that clonal identity is not necessarily inferred by acceptance of grafts. For M. dilatata, 18 of 40 cases of fusion were between dissimilar genotypes and for M. verrucosa, 3 of 7 cases. In one case, fusion occurred between genotypes which shared no alleles. A greater understanding of the genetics of invertebrate immunology is required before histocompatibility criteria can be used independently to assess population genetic structure with confidence.     

Experimental ecology of the temperate scleractinian coral Astrangia danae

The combined effects of temperature, light and symbiont density on the metabolic rate and calcification of the temperate coral Astrangia danae were studied experimentally using colonies containing different concentrations of zooxanthellae. After acclimation to five temperatures between 6.5° and 27°C, and incubation at three light levels and in darkness, respiration and photosynthesis were measured and corrected for rates due to commensals alone. Calcification rates were regressed on zooxanthellae concentration and production in order to define “symbiotic” and “non-symbiotic” averages, and the enhancement of calcification by symbiotic interactions in the polyps. Respiration by the polyparium varied less with temperature between 11.5° and 23°C than that of the commensals, suggesting physiological acclimation by the coral tissue. In-vivo zooxanthellae photosynthesis increased linearly with temperature and was near its maximum at 400 μEin m^?2 s^?1, but the photosynthesis of the endolithic algae of the corallum varied little between 11.5° and 27°C. Calcification at any given temperature was near its maximum at 40 μEin m^?2 s^?1 in both symbiotic and non-symbiotic corals. CaCO₃ deposition increased linearly with temperature in non-symbiotic colonies and in symbiotic colonies incubated in the dark. In symbiotic colonies, calcification in the light increased above these basic rates as temperature rose above 15°C. Below 15°C, symbiotic interactions failed to stimulate calcification, apparently due both to a lowering of zooxanthellae photosynthesis and to a decrease in the enhancing effect of any given level of primary production.     

Effect of light quality on photosynthesis of the reef coral Montipora verrucosa

Pieces of the reef coral Montipora verrucosa (Lam.), collected from Kaneohe Bay, Oahu, Hawaii in 1982, were grown in four low-light treatments (11% sunlight): blue, green, red and the full spectrum of photosynthetically active radiation (PAR); and at high-intensity full PAR (90% sunlight). These acclimated corals were then tested for photosynthetic ability in blue, green, red, and white light. The photosynthetic parameters that were measured were; ligh-saturated photosynthetic rate, the initial slope of the photosynthesis/irradiance curve, the light intensity where these two lines crossed, and dark respiration. While acclimation intensity had a pronounced effect, the results also showed that the color of the acclimation treatment influenced the photosynthetic responses of the corals. The color of the light used in the measurements of photosynthesis had much less effect on the photosynthetic responses of the corals.Contribution No. 729 of the Hawaii Institute of Marine Biology     

Oculina patagonica: a non-lessepsian scleractinian coral invading the Mediterranean Sea

The scleractinian coral Oculina patagonica De Angelis is a new immigrant from the Southwest Atlantic to the Mediterranean Sea, having established itself only recently along the Israeli coast. This species is the only scleractinian coral reported to have invaded a new region. In order to understand the swift establishment of this species along the Israeli coast, from 1994 to 1999 we studied its distribution, abundance, reproduction, recruitment, survival, and the effect of bleaching events on its population abundance. In addition, population studies of O. patagonica were performed at several localities along the eastern and western Mediterranean coasts. Highest abundance was recorded along the Spanish coast, reaching 30lj colonies per 10 m line transect in shallow water. Second in abundance was the Israeli coast, with a maximum of 10DŽ colonies per transect. O. patagonica was rare along the coast of Italy, and absent along the Mediterranean coast of France. During the study, both mortality and recruitment along the Israeli coast were very low. In contrast, recruitment along the Spanish coast was very high. Reproduction of the species was studied using gonadal histology. O. patagonica is gonochoric. Female gonads were first observed in May and male gonads in July, both reaching maturity in late August and early September. Naturally occurring azooxanthellate colonies of O. patagonica inhabiting small dark caves developed gonads and spawned in parallel to zooxanthellate colonies exposed to light. No gonads were found in zooxanthellate colonies that underwent bleaching during the reproduction season. The high incidence of bleaching events along the Israeli coast observed throughout the years of this study may explain the low recruitment of new colonies during the same period. In view of its current recruitment patterns, we expect further expansion of O. patagonica in range and abundance in the western Mediterranean, but very small expansion of the population in the eastern Mediterranean, due to repetitive annual bleaching events.     

Synergistic effects of temperature,salinity and light on the hermatypic coral Montipora verrucosa

Temperature tolerance in the reef coral Montipora verrucosa (Lamarck) is affected by salinity and light. Low salinity reduces ability of the coral to survive shortterm exposure to elevated temperature. High natural light intensity aggravates damage sustained by corals at high temperature. In long-term growth experiments, high light intensity caused substantial loss of zooxanthellar pigment, higher mortality rates, reduced carbon fixation and lowered growth rate at both upper and lower sublethal temperatures Effects of light at optimal temperature were less dramatic. Interactions between physical environmental factors appear to be most important near the limits of tolerance for a given factor. Acclimation capability was indicated, and was influenced by both thermal history and pigmentation state of stressed corals.Contribution No. 543 of the Hawaii Institute of Marine Biology.     

Cell architecture of the dinoflagellate Symbiodinium sp. inhabiting the Hawaiian stony coral Montipora verrucosa

The Hawaiian stony coral Montipora verrucosa (Lamarck) Quoy et Gaimard (Anthozoa) harbours zooxanthellae of the genus Symbiodinium Freudenthal (Dinophyceae). The algae occur as coccoid cells when inside their host and produce periodically motile flagellate cells when in culture. Coccoid cells of cultured specimens were investigated in this study, using three-dimensional reconstructions in tandem with quantitative analyses after electron microscopy of serially sectioned cells, as well as freeze-fracture electron microscopy. The amphiesma normally consists of five membraneous layers with intermediate material of unknown composition. The intracellular morphology is characterized by a single, peripheral, multilobed chloroplast with a parallel thylakoid arrangement, polyhedral inclusions resembling carboxysomes, and one double-stalked pyrenoid, outlined by a triple-layered chloroplast envelope. Spherical, elongated or branched mitochondria are aggregated in the center of the cell, surrounded by the chloroplast. The nucleus is a large, spherical structure located rather ventrally and containing 26 chromosomes with ovoid to elongated shapes. Further structures found to be present include Golgi apparatus, fibrous bodies, centrioles, and vacuoles containing crystals. Cell models of Symbiodinium sp. are represented in order to uncover completely the cellular microarchitecture of a gymnodinioid zooxanthella.     

Pink-line syndrome, a physiological crisis in the scleractinian coral Porites lutea

Coral diseases are one of the major factors that alter coral cover and their diversity. We have earlier reported the “Pink-line syndrome” (PLS) in the scleractinian coral Porites lutea wherein a colored band appears between the dead and healthy tissue of a colony. About 20% of the P. lutea colonies were affected in Kavaratti of the Lakshadweep Islands in the Arabian Sea during April 1996 and the incidence increased fourfold within the next 4 years. Fungi were associated in both PLS-affected and healthy specimens, whereas the cyanobacterium Phormidium valderianum occurred exclusively in the PLS-affected specimens. There was an increased expression of a 29 kDa protein without any significant increase in total protein content in the PLS-affected colonies. A reduced number of zooxanthellae and an increase in zooxanthellae size, mitotic index, and chl a concentrations were some of the characteristics of the PLS-affected colonies. PLS induction experiments conducted using selected fungi and the cyanobacterium P. valderianum isolated from the affected colonies and abiotic factors, such as CO₂ enrichment and the effect of cyanobacterial photosynthesis inhibition, indicated that the CO₂ build-up around the host tissue caused the pink coloration. We hypothesize that these physiological changes disturb the mutualism between the zooxanthellae and the host. When the symbiosis is disturbed by the external CO₂, the host loses control over the zooxanthellae, causing their uncontrolled division. This process may lead to a break in photosynthate transfer to the host, thereby resulting in starvation and finally leading to partial mortality. We further hypothesize that these degenerative processes are triggered by the CO₂ produced by P. valderianum through its carbon concentration mechanism. In this context, any opportunistic cyanobacteria or other agents having potential to interfere with the physiology of the host or the symbiont can cause such a physiological disorder. The mechanism of PLS formation is an early warning to protect corals as the increasing atmospheric CO₂ could induce PLS-like physiological disorder in corals.     

Reproductive periodicity of the scleractinian coral Lophelia pertusa from the Trondheim Fjord, Norway

Between 2002 and 2008, samples of the cold-water scleractinian coral Lophelia pertusa were collected from the Trondheim Fjord in Norway to examine reproductive periodicity. Collections were made from three locations: Tautra, (63°35.36′N, 10°31.23′E at 40–70 m), Stokkbergneset (63°28.18′N, 09°54.73′E at 110–500 m), and Røberg (63°28.88′N, 09°59.50′E at 250 m). Populations of L. pertusa from the Trondheim Fjord initiated oogenesis in January and spawning occurred from late January to early March the following year. Gametogenic cycles of the female L. pertusa samples overlapped by approximately 2 months, with oogonia visible in January, but this was not evident in the males. This paper provides the most complete gametogenic cycle to date and spawning observations for this important structure-forming species. The results from fjord populations are compared with published and preliminary data from other regions and are discussed in the context of regional differences in physical and biological variables, particularly food supply. Differences in gametogenic cycles within a single species provide a rare opportunity (especially in deep-sea species) to examine potential drivers of reproduction.     

Using three-dimensional surface area to compare the growth of two Pocilloporid coral species

Many facets of coral research require coral colony surface area estimates. This study developed a relationship between the two-dimensional (2D) projected area and the three-dimensional (3D) whole colony surface area for two commonly studied Indo-Pacific coral species: Pocillopora damicornis and Stylophora pistillata. The surface index function was used to measure the growth of colonies in situ around Heron reef on the southern Great Barrier Reef. The results show that while growth between the two species was not significantly different when measured in two dimensions, the 3D area showed significantly different growth rates with S. pistillata growing at almost double the rate of P. damicornis. The study demonstrates that it is possible to make reliable estimates of the 3D surface area of entire colonies of these complex branching coral species, using the plan view of the coral and a pre-determined surface index function. In addition, this study shows that the 3D surface area provides a more useful measure of colony growth than the traditional methods of either 2D area or longest dimension.     

Evidence for broadcast spawning as well as brooding in the scleractinian coral Pocillopora damicornis

The reproductive activity of 88 colonies of Pocillopora damicornis at Rottnest Island, Western Australia, was examined over 12 mo (20 August 1988 to 20 August 1989). Larvae were found close to the three new moons of February, March and April, 1989, but in large numbers only during March; 67% of all corals produced larvae. There was no obvious connection between gametogenesis and the production of brooded larvae; however, those corals which did not produce, larvae also did not produce ova but did produce, sperm, and must be considered male colonies. All colonies which produced ova went on to produce planulae. Spermatogenesis occurred throughout the year, but oogenesis occurred between September 1988 and April 1989. Dramatic declines in the percentage of polyps containing eggs and sperm occurred in early February and early April 1989. On these dates the amount of lipid in the tissue of the corals decreased, which suggests that the decline in the percentage of polyps with sperm and eggs was not due to the gametes being resorbed by the polyps. Therefore, P. damicornis at Rottnest Island appears to be both a brooder and a broadcaster with asexual reproduction through brooded larvae and sexual reproduction through spawning of gametes. This accounts for earlier evidence in the literature that some input from sexual reproduction had been occurring in the populations of P. damicornis at Rottnest Island. There is no direct evidence for spawning, as broadcasting of gametes has not been observed, but the indirect evidence provides a strong argument for the existence of this dual reproduction.     

Allozyme and colour differences between two sibling species of the heteronemertean Lineus torquatus from the Sea of Japan

Brown and cherry colour morphs of the heteronemertean Lineus torquatus Coe, 1901 from the Vostok Bay of the Sea of Japan were compared using allele frequencies at 18 isozyme loci. These morphs shared no common alleles at 9 of the 18 loci. The observed genetic differences are strong evidence that the brown and cherry colour morphs, which occur sympatrically, are different species. Nei's genetic distance between these species (D=1.135) is considerably higher than typical D values for congeneric animal species. As with much previous work on nemerteans, intraspecific allozymic variability in both species was high, with mean expected heterozygosity values 0.286±0.049 and 0.159±0.051 for L. torquatus (brown) and L. torquatus (cherry) species, respectively.     

Effects of ambient levels of solar ultraviolet radiation on zooxanthellae and photosynthesis of the reef coral Montipora verrucosa

Paired flat plates of the hermatypic coral Montipora verrucosa from Kaneohe Bay, Oahu, Hawaii, were acclimated to photosynthetically active radiation (PAR) only and to full sunlight (PAR+UV) for several weeks in the summer of 1990. After the acclimation period, photosynthesis, both in PAR-only and PAR+UV as well as dark respiration were measured. Levels of the UV-absorbing compounds, S320, density of zooxanthellae, and chlorophyll a concentration were determined. Corals acclimated in PAR+UV had higher levels of the UV-protective compounds and lower areal zooxanthellae densities than corals acclimated in PAR-only. Chlorophyll a per unit volume of coral host and per algal cell did not differ between corals from the two acclimation treatments. Corals acclimated to PAR+UV displayed higher photosynthesis in full sunlight than corals acclimated to PAR-only, but when photosynthesis was measured in the light regime to which the corals had been acclimated, there were no differences in photosynthesis. Dark respiration was the same for corals from the two acclimation treatments regardless of the light quality immediately preceding the dark period.Contribution No. 902 HIMB     

Substratum preferences in planula larvae of two species of scleractinian corals, Goniastrea retiformis and Stylaraea punctata

To test whether coral planulae recruit randomly to different coral reef habitats or have specific substratum preferences, the settling behavior of planulae from two shallow water coral species from Pago Bay, Guam (13°25.02N, 144°47.30E) were examined in the laboratory in June and July of 1995. Goniastrea retiformis is generally restricted to the shallow reef front (<10 m depth) in areas dominated by crustose coralline algae (CCA), while Stylaraea punctata is abundant on inner reef flats were CCA coverage is low and sand and carbonate rubble covered by biofilms is common. When presented with four substrata (1) carbonate rock scrubbed free of biofilm and dried as a control, (2) the CCA Hydrolithon reinboldii, (3) the CCA Peyssonelia sp., and (4) naturally conditioned carbonate rubble covered by a biofilm, G. retiformis larvae showed a significant preference for H. reinboldii, and S. punctata larvae for the carbonate biofilm treatment. The preference shown by S. punctata larvae for biofilmed surfaces did not diminish with increasing larval age up to 11 days. These results suggest that the larvae of both species are capable of habitat selection, and that the preferred substrata among those tested bears a relationship to the habitats in which adult colonies were found. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Burrowing by the bivalve mollusc Lithophaga curta in the living reef coral Montipora berryi and a hypothesis of reciprocal larval recruitment

The burrowing bivalve Lithophaga curta is abundant (8 to 9 per 100 cm²) in the encrusting, hermatypic coral Montipora berryi at Enewetak; this is the first report of L. curta colonizing living coral. Conversely, larvae of M. berryi appear to settle in empty bivalve burrows, tentatively identified as those of L. curta, located in dead M. berryi. Several hypotheses are suggested on the adaptive significance of substrate selection by larvae of both species and an overall hypothesis of reciprocal recruitment, describing the dynamic aspects of this apparently co-evolved relationship, is proposed.