δ<Superscript>13</Superscript>C and δ<Superscript>15</Superscript>N values in reef corals <Emphasis Type="Italic">Porites lutea</Emphasis> and <Emphasis Type="Italic">P. cylindrica</Emphasis> and in their epilithic and endolithic algae

In summer 1998, shallow water corals at Sesoko Island, Japan (26°38′N, 127°52′E) were damaged by bleaching. In August 2003, partially damaged colonies of the massive Porites lutea and the branching P. cylindrica were collected at depths of 1.0–2.5 m. The species composition of epilithic algal communities on dead skeletal surfaces of the colonies (‘red turfs’, ‘green turfs’, ‘red crusts’) and the endolithic algae (living in coral skeletons) growing close to and away from living coral polyps was determined. Carbon and nitrogen stable isotope values of organic matter (δ¹³C and δ¹⁵N) from all six of these biological entities were determined. There were no significant differences in the isotope composition of coral tissues of the two corals, with P. lutea having δ¹³C of −15.3 to −9.6‰ and δ¹⁵N of 4.7–6.1‰ and P. cylindrica having similar values. Polyps in both species living close to an interface with epilithic algae had similar isotope values to polyps distant from such an interface. Despite differences in the relative abundance of the algal species in red turfs and crusts, their δ¹³C and δ¹⁵N values were not significantly different from each other (−18.2 to −13.9, −20.6 to −16.2, 1.1–4.3, and 3.3 to 4.9‰, respectively). The green algal turf had significantly higher δ¹³C values (−14.9 to −9.3‰) than that of red turfs and crusts but similar δ¹⁵N (1.2–4.1‰) to the red algae. The data do not suggest that adjoining associations of epilithic algae and coral polyps exchange carbon- and nitrogen-containing metabolites to a significant extent. The endolithic algae in the coral skeletons had δ¹³C values of −14.8 to −12.3‰ and δ¹⁵N of 4.0–5.4‰. Thus they did not differ significantly from the coral polyps in their carbon and nitrogen isotope values. The similarity in carbon isotope values between the coral polyps and endolithic algae may be attributed to a common source of CO₂ for zooxanthellae and endolithic algae, namely, from respiration by the coral host. While it is difficult to fully interpret similarity in the nitrogen isotope composition of coral tissue and of green endolithic algae and the difference in δ¹⁵N between green epilithic and endolithic algae, the data are consistent with nitrogen-containing metabolites from the scleractinian coral serving as a significant source of nitrogen for the endolithic algae.     

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     

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.     

Effects of dissolved ammonium addition and host feeding with Artemia salina on photoacclimation of the hermatypic coral Stylophora pistillata

 Effects of nutrient treatments on photoacclimation of the hermatypic coral Stylophora pistillata (Esper) were studied. Studies on photoacclimation of colonies from different light regimes in the field were evaluated and used to design laboratory experiments. Coral colonies were collected in the Gulf of Eilat (Israel) from January to March 1993. Exterior branches of colonies from different depths (1 to 40 m) displayed different trends in production characteristics at reduced and very low levels of illumination. From 24 ± 3% to 12 ± 2% of incident surface photosynthetic active radiation (PAR_o), zooxanthella population density and chlorophyll a+c per 10⁶ zooxanthellae increased, a trend seen in the range of light levels optimal for coral growth (90 to 30% PAR_o). The P _max of CO₂ per 10⁶ zooxanthellae decreased, while P _max of CO₂ per 10³ polyps increased, indicating an increase in zooxanthella population density at low light levels. Proliferous zooxanthella frequency (PZF, a measure of zooxanthella division) declined significantly at light levels <18 ± 3% PAR_o. At the lowest levels of illumination (<5% PAR_o), zooxanthella population density decreased, as did the PZF; chl a+c per 10⁶ zooxanthellae was unchanged. In 28-d experiments, exterior coral branches from the upper surfaces of colonies from 3 m depth (65 ± 4% PAR_o) were incubated in aquaria under bright (80 to 90% PAR_o), reduced (20 to 30% PAR_o), and extremely low (2 to 4% PAR_o) light intensities. At each light intensity, the corals were maintained in three feeding treatments: sea water (SW); ammonium enriched SW (SW + N); SW with Artemia salina nauplii (SW + A). An increase in P _max of CO₂ per 10³ polyps was found in corals acclimated to reduced light (20 to 30% PAR_o) in nutrient-enriched SW, while in SW, where the increase in zooxanthella population density was smaller, it did not occur. Nutrient enrichments (SW + N at 2 to 4% PAR_o and SW + A at 20 to 30% PAR_o) increased zooxanthella population density, but had no effect on chl a+c per 10⁶ zooxanthellae. Acclimation for 14 d to reduced (10 to 20% PAR_o) and extremely low (1 to 3% PAR_o) light intensities shifted ¹⁴C photoassimilation into glycerol and other compounds (probably glycerides), rather than sugars. Both ammonium addition and feeding with Artemia salina nauplii resulted in an increase in photosynthetic assimilation of ¹⁴C into amino acids. We conclude that acclimation to reduced light consists of two processes: an increase in photosynthetic pigments and in zooxanthella population density. Both processes require nitrogen, the increase in zooxanthella population density needing more; this adaptation is therefore limited in nitrogen-poor sea water. Received: 19 June 1998 / Accepted: 13 June 2000     

Experimental ecology of the temperate scleractinian coral Astrangia danae I. Partition of respiration,photosynthesis and calcification between host and symbionts

Calcification, photosynthesis and respiration of the scleractinian coral Astrangia danae were calculated from the changes in total alkalinity, pH, calculated total CO₂, and oxygen concentration produced by colonies incubated in glass jars. A correction for changes in ammonia, nitrate and nitrite was taken into account and the method evaluated. The fluxes of oxygen and CO₂ were highly correlated (r=0.99). The statistical error of alkalinity determinations was less than 10% of the changes observed in the slowest calcifying samples. Metabolism of polyparium alone was estimated by difference after removal of tissue and reincubation of bare corallum. Zooxanthellae concentration in the polyps was obtained from cell counts made on homogenates of polyp tissue. The calculated photosynthetic rate of the zooxanthellae in vivo was 25 mol O₂ (10⁸ cell)^-1 h^-1 at a light intensity of 120 Ein m^-2 s^-1. In corals having 0.5x10⁹ zooxanthellae/dm² of colony area up to 8% of the total photosynthesis was attributed to the corallum microcosm. Polyp respiration, photosynthesis, and CaCO₃ uptake rates were all much higher than rates previously reported from A. danae, apparently because in these experiments the organisms were better fed. This increased photosynthesis in turn enhanced calcification still further. The symbiosis therefore appears to provide a growth advantage even to fed corals, under the conditions of these experiments.     

Effects of feeding frequency and symbiosis with zooxanthellae on nitrogen metabolism and respiration of the coral Astrangia danae

Colonies of the temperate coral Astrangia danae occur naturally with and without zooxanthellae. Basal nitrogen excretion rates of nonsymbiotic colonies increased with increasing feeding frequency [average excretion rate was 635 ng-at N (mg-at tissue-N)^-1 h^-1]. Reduced excretion rates of symbiotic colonies were attributed to N uptake by the zooxanthellae. Nitrogen uptake rates of the zooxanthellae averaged 8 ng-at N (10⁶ cells)^-1 h^-1 in the dark and 21 ng-at N (10⁶ cells)^-1 h^-1 at 200 Ein m^-2 s^-1. At these rates the zooxanthellae could provide 54% of the daily basal N requirement of the coral if all of the recycled N was translocated. Basal respiration rates were 172 nmol O₂ cm^-2 h^-1 for starved colonies and 447 nmol O₂ cm^-2 h^-1 for colonies fed three times per week. There were no significant differences between respiration rates of symbiotic and nonsymbiotic colonies. N excretion and respiration rates of fed (symbiotic and nonsymbiotic) colonies increased greatly soon after feeding. N absorption efficiencies decreased with increasing feeding frequency. A N mass balance, constructed for hypothetical situations of nonsymbiotic and symbiotic (3×10⁶ zooxanthellae cm^-2) colonies, starved and fed 15 g-at N cm^-2wk^-1, showed that the presence of symbionts could double the N growth rate of feeding colonies, and reduce the turnover-time of starved ones, but could not provide all of the N requirements of starved colonies. Rates of secondary production, estimated from rates of photosynthesis and respiration were similar to those estimated for reef corals.     

Do corals select zooxanthellae by alternative discharge?

Loss of zooxanthellae (dinoflagellate Symbiodinium) from corals will sometimes lead to mass mortality of corals. To detect and quantify Symbiodinium released from corals, we developed a zooxanthellae “trap” and a quantitative PCR (qPCR) system with Symbiodinium clades A–F-specific primer sets. The trap was attached to a branch or the surface of several wild stony corals, and the water samples within the traps, including released Symbiodinium, were subjected to qPCR. All tested corals released clade C Symbiodinium at estimates of ~5,900 cells h⁻¹ cm⁻² of coral surface. Although all tested Pocillopora eydouxi harboured both clades C and D, some of these colonies released only clade C or released a lesser amount of clade D than that in the tissues. Our Symbiodinium quantification system revealed that wild hermatypic corals constantly release Symbiodinium to the environment. Our result suggests that some corals may discharge certain clades of Symbiodinium alternatively.     

A “CO2 supply” mechanism in zooxanthellate cnidarians: role of carbonic anhydrase

Carbonic anhydrase (CA, EC 4.2.1.1) activity was detected in 22 species of tropical cnidarians which contain endosymbiotic dinoflagellates (=zooxanthellae). CA activity was 2 to 3 times higher in animal tissue than in algae and ca. 29 times higher in zooxanthellate than azooxanthellate species. It was also higher in the zooxanthellate tentacle tissue than in the azooxanthellate column tissue of the anemone Condylactis gigantea. CA was therefore significantly related to the presence of endosymbiotic algae. Further results indicated that CA functions in the photosynthetic carbon metabolism of zooxanthellate cnidarians as evidenced by (1) low CA activity in shade-adapted and deep water colonies compared to the more productive shallow water, light-adapted colonies of the coral Stylophora pistillata, and (2) the 56 to 85% reduction in photosynthetic carbon assimilation by zooxanthellae in situ in the presence of Diamox, an inhibitor of CA. Although CA has been proposed to function in calcification, its association with zooxanthellae and photosynthetic activity in both calcifying and non-calcifying associations suggests a role in photosynthetic metabolism of algal/cnidarian symbioses. It is proposed that CA acts as a CO₂ supply mechanism by releasing CO₂ from bicarbonate, and enabling zooxanthellae to maintain high rates of photosynthesis in their intracellular environment.     

Three party symbiosis: acoelomorph worms,corals and unicellular algal symbionts in Eilat (Red Sea)

Epizoic worms were found to occur on certain coral colonies from reefs off the coast of Eilat (Red Sea). We identified 14 coral species infested by acoelomorph worms at a depth range of 2–50 m. The host corals were all zooxanthellate and included both massive and branching stony corals and a soft coral. Worms from all hosts were identified as belonging to the genus Waminoa and contained two distinct algal symbionts differing in size. The smaller one was identified as Symbiodinium sp. and the larger one is presumed to belong to the genus Amphidinium. Worm-infested colonies of the soft coral, Stereonephthya cundabiluensis, lacked a mucus layer and exhibited distinct cell microvilli, a phenotype not present in colonies lacking Waminoa sp. In most cases, both cnidarian and Acoelomorph hosts displayed high specificity for genetically distinctive Symbiodinium spp. These observations show that the epizoic worms do not acquire their symbionts from the “host” coral.     

High genetic diversity of the symbiotic dinoflagellates in the coral Pocillopora meandrina from the South Pacific

Symbioses between dinoflagellates in the genus Symbiodinium (commonly referred to as zooxanthellae) and scleractinian corals are an essential feature for the maintenance of coral reefs. The fine-scale diversity and population structure of the zooxanthellae inhabiting the coral Pocillopora meandrina, a major reef building species in Polynesia, was examined. We used two polymorphic microsatellites to study seven populations from the South Pacific, whose host structuring has been previously investigated. The symbionts of P. meandrina showed high levels of diversity, with more than one zooxanthella genotype being identified in most of the host individuals. Genetic differentiation between symbiont populations was detected at a large scale (2,000 km) between the Tonga and the Society Archipelagos. Within the Society Archipelago, the two most remote populations (Tahiti and Bora-Bora; 200 km apart) were only weakly differentiated from each other. Statistical tests demonstrated that the symbiont genetic structure was not correlated with that of its host, suggesting that dispersal of the symbionts, whether they are transported within a host larva or free in the water, depends mainly on distance and water currents. In addition, the data suggests that hosts may acquire new symbionts after maternal transmission, possibly following a disturbance event. Lastly, the weak differentiation between symbiont populations of P. verrucosa and P. meandrina, both from Moorea, indicated that there was some host-symbiont fine-scale specificity detectable at the genetic resolution offered by microsatellites.     

Adaptation strategies of the corallimorpharian <Emphasis Type="Italic">Rhodactis rhodostoma</Emphasis> to irradiance and temperature

Corallimorpharians may dominate some habitats on coral reefs and compete with stony corals for access to light, yet little is known concerning their photosynthetic traits. At Eilat in the northern Red Sea, we observed that the abundance of individuals of the corallimorpharian Rhodactis rhodostoma decreased significantly with depth on the reef slope. Field and laboratory experiments revealed that they employ several mechanisms of photoadaptation to high irradiance on the shallow reef flat. Their endosymbiotic microalgae (zooxanthellae) varied significantly in both abundance and chlorophyll content with level of irradiance. Use of a diving pulse amplitude modulated fluorometer revealed that the zooxanthellae of R. rhodostoma effectively disperse excess light energy by expressing significantly higher values of non-photochemical quenching and maximum excitation pressure on photosystem II when experimentally exposed to high light (HL) versus low light (LL). Host corallimorpharian tissues mediated this response by shielding the algal symbionts from high irradiance. The endoderm of host tentacles thickened significantly and microalgal cells were located further from the mesoglea in HL than in LL. The clades of zooxanthellae hosted by the corallimorpharians also varied with depth. In shallow water, all sampled individuals hosted clade C zooxanthellae, while in deep water the majority hosted clade D. The photosynthetic output of individuals of R. rhodostoma was less affected by HL than was that of a stony coral examined. When exposed to both high temperature (HT) and HL, individuals of R. rhodostoma reduced their maximum quantum yield, but not when exposed to HL at low temperature (LT). In contrast, colonies of the scleractinian coral Favia favus reduced their photosynthetic output when exposed to HL in both temperature regimes. After 2 weeks of HT stress, R. rhodostoma polyps appeared to bleach completely but re-established their zooxanthella populations upon return to ambient temperature. We conclude that mechanisms of photoadaptation to high irradiance employed by both the endosymbiotic zooxanthellae and host corallimorpharians may explain in part the abundance of R. rhodostoma on some shallow reef flats. The ability to survive for weeks at HT while bleached also may allow corallimorpharians to repopulate shallow reef areas where scleractinians have been killed by thermal stress. B. Kuguru and G. Winters contributed equally to this work.     

Adaptation of solitary corals and their zooxanthellae to low light and UV radiation

Adaptation of solitary corals, Fungia repanda and F. echinata, and their zooxanthellae to low light and ultraviolet light B (UV-B) was studied with respect to changes in their protein contents, photosynthetic pigment contents and the photosynthesis-irradiance (P-I) curves. The corals were collected from 1 to 50 m depths in the Republic of Belau (Paulau) in 1990 and 1991. The chlorophyll a content in a unit surface area of the coral did not change significantly with the depth of the habitat, whereas cellular chlorophyll a in the algae increased with the depth. Zooxanthellae density and protein content in a unit surface area of Fungia spp. decreased with the depth. Photosynthetic parameters normalized by a unit surface area of the Fungia spp., maximum gross photosynthetic rate (P _gmax area^-1) and dark respiration rate (R area^-1), were negatively correlated with the depth, while initial slope of the P-I curve () did not show significant correlation with the depth. Compensation light intensity (Ic) decreased with the depth. In isolated zooxanthellae, P _max chl a ^-1, and R chl a ^-1 decreased with the depth, while _{chl a} was constant. P _gmax cell^-1 and R cell^-1 did not change significantly but _cell increased with the depth. Ic decreased with the depth as in the intact corals. Reduction of protein content in a unit area of the coral from deeper habitat implies decrease of host animal tissues. Reduction of Ic can be explained by decrease of R area^-1, which may be due to the diminution of animal tissues. The photoadaptational response to low light intensity of intact Fungia spp. was found to be a combination of the photoadaptation of symbiotic algae and the decrease of host animal tissue. In order to study their adaptation to ultraviolet (UV) radiation, P-I curves of Fungia spp. and isolated zooxanthellae were analyzed before and after UV-B irradiation. 1 h UV-B irradiation showed no effect on the photosynthetic rate of the shallow water (1 m) corals, while it inhibited the photosynthesis of the deep water (30 m) corals and zooxanthellae isolated from both shallow and deep water corals. These results indicate that the host, Fungia spp., in shallow water have protective mechanism for intense UV-B in their habitat. These photoadaptational mechanisms seem to allow the Fungia spp. to have wide vertical distribution where light intensity spans more than two orders of magnitude.     

Stimulation of fat-body production in the polyps of the coralPocillopora damicornis by the presence of mutualistic crabs of the genusTrapezia

A mutualism exists between the xanthid crabs of the genusTrapezia and their host corals,Pocillopora damicornis. It has previously been established that these obligate coral residents benefit the coral hosts by defending them against echinoderm predators and by increasing the survival of polyps located deep between the coral branches. In turn, the corals apparently benefit the crabs by producing lipid-filled structures on which the trapezid crabs feed; these fat bodies may contain some of the lipid which in previous studies of coral metabolism has been termed excess. It was determined by experiments conducted at the Hawaii Institute of Marine Biology that the presence of crabs in colonies ofP. damicornis stimulates the polyps to produce the lipid-filled fat bodies; removal of crabs causes corals to cease producing fat bodies. A structure very similar to the fat bodies ofP. damicornis has been reported inAcropora durvillei. Both of these coral genera ordinarily possess xanthid-crab mutualists. This association between branching corals and crustaceans may have evolved because corals of these genera provide shelter among their branches and because these shallow-water corals are evidently capable of releasing lipid which is excess to the corals' metabolic needs, but which can be utilized by the crabs.     

Effects of nutritional history on nitrogen assimilation in congeneric temperate and tropical scleractinian corals

The nutritional history of corals is known to affect metabolic processes such as inorganic nutrient uptake and photosynthesis, but little is known about how it affects assimilation efficiency of ingested prey items or the partitioning of prey nitrogen between the host and symbiont. The temperate scleractinian coral Oculina arbuscula and its tropical congener Oculina diffusa were acclimated to three nutritional regimes (fed twice weekly, starved, starved with an inorganic nutrient supplement), then fed Artemia nauplii labeled with the stable isotope tracer ¹⁵N. Fed corals of both species had the lowest assimilation efficiencies (36–51% for O. arbuscula, 38–57% for O. diffusa), but were not statistically different from the other nutritional regimes. Fed and starved corals also had similar NH₄⁺ excretion rates. This is inconsistent with decreased nitrogen excretion and reduced amino acid catabolism predicted by both the nitrogen recycling and conservation paradigms. In coral host tissue, ~90% of the ingested ¹⁵N was in the TCA-insoluble (protein and nucleic acids) and ethanol-soluble (amino acids/low molecular weight compounds) within 4 h of feeding. The TCA-insoluble pool was also the dominant repository of the label in zooxanthellae of both species (40–53% in O. arbuscula, 50–60% in O. diffusa). However, nutritional history had no effect on the distribution of prey ¹⁵N within the biochemical pools of the host or the zooxanthellae for either species. This result is consistent with the nitrogen conservation hypothesis, as preferential carbon metabolism would minimize the effects of starvation on nitrogen-containing biochemical pools.Communicated by P.W. Sammarco, Chauvin     

Effects of sheltering fish on growth of their host corals

Stony corals are the foundation species of tropical reefs, and their structures can harbor a diverse range of mutualist taxa that can confer important benefits, including provision of nutrients. Prominent among the associates of branching coral in the genus Pocillopora are groups of zooplanktivorous damselfishes that take refuge in the coral to avoid their predators. In field and laboratory experiments, we explored the effects of colonies of resident damselfishes on growth of their host corals. Laboratory studies revealed a positive relationship between biomass of fish and output of ammonium. In the field, levels of ammonium were significantly elevated in the water surrounding the branches of Pocillopora occupied by colonies of damselfish, particularly in time periods following active feeding by the fish. Experimental manipulation of the presence of fish on host corals during a month-long field experiment revealed that corals hosting fish grew significantly more than those that lacked fish, and coral growth was positively correlated with the biomass of resident fish. The Pocillopora colonies in the field experiment varied in the degree of openness of their branching structure, and dye studies indicated that this affected their ability to retain waterborne nutrients. Together with biomass of resident fish, colony openness explained 76% of the variation in coral growth rate during the experiment. Corals can exhibit considerable morphological variability, and mutualistic fish respond to colony architecture during habitat selection, with some species preferring more open-branched forms. This makes it likely that corals may face tradeoffs in attracting resident fish and in retaining the nutrients they provide.     

Effects of pCO2 on the interaction between an excavating sponge,Cliona varians,and a hermatypic coral,Porites furcata

Rising dissolved pCO₂ is a mounting threat to coral reef ecosystems. While the biological and physiological impacts of increased pCO₂ are well documented for many hermatypic corals, the potential effects on bioerosion processes remain largely unknown. Increases in pCO₂ are likely to modify the direct interactions between corals and bioeroders, such as excavating sponges, with broad implications for the balance between biologically mediated deposition and erosion of carbonate in reef communities. This study investigated the effects of three levels of CO₂ (present-day, mid-century and end-of-century projections) on the direct interaction between a bioeroding sponge, Cliona varians, and a Caribbean coral, Porites furcata. Increased pCO₂ concentrations had no effect on the attachment rates of C. varians to the corals, and we observed no significant impact of pCO₂ on the survival of either the coral or sponges. However, exposure to end-of-century levels of CO₂-dosing (~750 μatm) reduced calcification in P. furcata and led to a significant increase in sponge-mediated erosion of P. furcata. These findings demonstrate that pCO₂ can enhance erosional efficiency without impacting survival or competitive vigor in these two species. While few studies have considered the influence of pCO₂ on the competitive outcomes of interactions between corals and other reef organisms, our study suggests that assessing the impacts of changing pCO₂ on species interactions is crucial to adequately predict ecosystem-level responses in the future.     

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     

Microbial consortia increase thermal tolerance of corals

This study examined the response of a coral holobiont to thermal stress when the bacterial community was treated with antibiotics. Colonies of Pocillopora damicornis were exposed to broad and narrow-spectrum antibiotics targeting coral-associated α and γ-Proteobacteria. Corals were gradually heated from the control temperature of 26 to 31 °C, and measurements were made of host, zooxanthellar and microbial condition. Antibiotics artificially reduced the abundance and activity of bacteria, but had minimal effect on zooxanthellae photosynthetic efficiency or host tissue protein content. Heated corals without antibiotics showed significant declines in F _V /F _M , typical of thermal stress. However, heated corals treated with antibiotics showed severe tissue loss in addition to a decline in F _V /F _M . This study demonstrated that a disruption to the microbial consortium diminished the resilience of the holobiont. Corals exposed to antibiotics under control temperature did not bleach, suggesting that temperature may be an important factor influencing the activity, diversity and ecological function of the holobiont bacterial community.     

Depth-dependent responses to solar ultraviolet radiation and oxidative stress in the zooxanthellate coral Acropora microphthalma

Colonies of Acropora microphthalma (Verrill 1869) were transferred from depths of 2 to 3, 10, 20, and 30 m to UV-transparent and UV-opaque respirometry chambers placed at 1 m depth at Bowl Reef, Great Barrier Reef, in March 1989. Peak rates of photosynthesis in colonies originating at 2 and 10 m were unaffected by solar ultraviolet (UV) radiation at 1 m, whereas photosynthesis showed 30 and 38% inhibition in colonies transferred from 20 and 30 m, respectively. This differential sensitivity of corals to UV radiation was consistent with the five- to tenfold higher concentrations of UV-absorbing, mycosporine-like amino acids (MAAs, putative defenses against UV) in 2- and 10-m colonies compared with 20- and 30-m colonies. Photosynthesis in zooxanthellae freshly isolated from 2- and 10-m corals, however, was inhibited by UV, indicating that the host's tissues, which contain 95% of the total MAAs in corals at these depths, are the first line of defense against solar UV and provide protection to their endosymbiotic algae. The general bathymetric decline in the activities of the antioxidant enzyme superoxide dismutase (SOD) in the host, and SOD, catalase, and ascorbate peroxidase in the zooxanthellae, is related to the decrease in potential for photooxidative stress with increasing depth.     

Lipogenesis in the intact coral Pocillopora capitata and its isolated zooxanthellae: Evidence for a light-driven carbon cycle between symbiont and host

Surface tissue of the reef coral Pocillopora capitata contained approximately 34% lipid on a dry weight basis. Of this, 75% was storage lipid (wax ester and triglyceride) and 25% structural (phospholipid, galactolipid, etc.). Based on chlorophyll a: lipid ratios of intact coral and isolated zooxanthellae, it was determined that over 90% of the storage lipid resided in the host tissue. One half of the structural lipids was found in the host and the other in the symbiotic algae. Gentle fractionation of coral tissue indicated that zooxanthellae possessed less than 14% of the total coral protein. Coral tips and isolated zooxanthellae were incubated with sodium acetate-1-¹⁴C in light and dark to obtain lipogenic rates and proportions of fatty acids and lipid classes synthesized. The rate of lipid synthesis from acetate-1-¹⁴C by intact coral was stimulated three-fold in the light (1200 lux), which indicated that the majority of coral lipogenesis occurred in the zooxanthellae. Intact coral triglycerides contained ca. 68% of the ¹⁴C-activity and wax esters ca. 21%. Zooxanthellae isolated by the Water Pik technique synthesized negligible amounts of wax ester, which implied that wax ester synthesis was a property of the animal tissue. Isolated zooxanthellae and intact coral synthesized identical triglyceride fatty acids from acetate-1-¹⁴C. This study provides evidence for a carbon cycle between host and symbiont whereby the zooxanthellae take up host-derived carbon (probably in the form of acetate), synthesize fatty acids using their photosynthetically derived energy, and return the lipid to the host where it appears as wax ester and triglyceride.