Caribbean massive corals not recovering from repeated thermal stress events during 2005–2013

Massive coral bleaching events associated with high sea surface temperatures are forecast to become more frequent and severe in the future due to climate change. Monitoring colony recovery from bleaching disturbances over multiyear time frames is important for improving predictions of future coral community changes. However, there are currently few multiyear studies describing long‐term outcomes for coral colonies following acute bleaching events. We recorded colony pigmentation and size for bleached and unbleached groups of co‐located conspecifics of three major reef‐building scleractinian corals (Orbicella franksi, Siderastrea siderea, and Stephanocoenia michelini; n = 198 total) in Bocas del Toro, Panama, during the major 2005 bleaching event and then monitored pigmentation status and changes live tissue colony size for 8 years (2005–2013). Corals that were bleached in 2005 demonstrated markedly different response trajectories compared to unbleached colony groups, with extensive live tissue loss for bleached corals of all species following bleaching, with mean live tissue losses per colony 9 months postbleaching of 26.2% (±5.4 SE) for O. franksi, 35.7% (±4.7 SE) for S. michelini, and 11.2% (±3.9 SE) for S. siderea. Two species, O. franksi and S. michelini, later recovered to net positive growth, which continued until a second thermal stress event in 2010. Following this event, all species again lost tissue, with previously unbleached colony species groups experiencing greater declines than conspecific sample groups, which were previously bleached, indicating a possible positive acclimative response. However, despite this beneficial effect for previously bleached corals, all groups experienced substantial net tissue loss between 2005 and 2013, indicating that many important Caribbean reef‐building corals will likely suffer continued tissue loss and may be unable to maintain current benthic coverage when faced with future thermal stress forecast for the region, even with potential benefits from bleaching‐related acclimation.     

Continuation of sexual reproduction in <Emphasis Type="Italic">Montipora capitata</Emphasis> following bleaching

Bleaching is generally expected to produce detrimental impacts on coral reproduction. This study compared the fecundity of bleached and unbleached colonies of the Hawaiian coral Montipora capitata. It was hypothesized that bleaching would have no effect on reproduction because previous studies have shown that Montipora capitata can increase heterotrophic feeding following bleaching. Reproductive parameters, total reproductive output (bundles released ml⁻¹ coral colony), number of eggs bundle⁻¹, and egg size, measured in the summer of 2005 did not differ between colonies that bleached or did not bleach during 2004. These data were collected following a single bleaching event and cannot be used to predict the outcome should bleaching episodes become more frequent or severe.     

Impacts of bleaching on the soft coral Lobophytum compactum. II. Biochemical changes in adults and their eggs

Experimental bleaching reduces the levels of important biochemical parameters in adult tissues and eggs of the soft coral Lobophytum compactum. Protein, lipid, mycosporine-like amino acids (MAAs) and carotenoid concentrations remained lower in bleached adults than in controls for at least 8 months. Reductions in concentrations of all four parameters were greater in eggs than in maternal tissues, potentially jeopardizing egg and larval viability. In particular, reductions in lipids, proteins and carotenoids in tissues of heavily bleached soft corals were amplified approximately twofold in eggs. In comparison, amplification of maternal tissue reductions were not as great for MAAs, suggesting that MAAs are given higher priority in egg provisioning. Our finding that MAA levels are normally three times higher in eggs than in unbleached maternal tissues supports the importance of MAAs for larval survival. Twenty months after experimental bleaching the biochemical composition of both adult tissues and their eggs were indistinguishable from those of control (unbleached) soft corals. Accepted: 1 June 2000     

Fast repetition rate (FRR) fluorometry: variability of chlorophyll a fluorescence yields in colonies of the corals, Montastraea faveolata (w.) and Diploria labyrinthiformes (h.) recovering from bleaching

Recently, an underwater version of a fast repetition rate fluorometer (FRRF) was developed for the non-destructive study of fluorescence yields in benthic photoautotrophs. We used an FRRF to study bleached colonies of the corals, Montastraea faveolata and Diploria labyrinthiformes at sites surrounding Lee Stocking Island, Exuma, Bahamas, to assess their recovery from bleaching ( approximately 1 year after the initial bleaching event) induced by elevated temperatures. The steady state quantum yields of chlorophyll a fluorescence (DeltaF'/F'(m)) from photosystem II (PSII) within coral colonies were separated into three categories representing visibly distinct degrees of bleaching ranging from no bleaching to completely bleached areas. Differences in DeltaF'/F'(m) were significantly different from bleached to unbleached regions within colonies. Dark, unbleached regions within colonies exhibited significantly higher DeltaF'/F'(m) values (0.438+/-0.019; mean+/-S.D.) when compared to lighter regions, and occupied a majority of the colonies' surface area (46-73%). Bleached regions exhibited significantly lower DeltaF'/F'(m) (0.337+/-0.014) and covered only 7-25% of the colonies' surface area. The observations from this study suggest that zooxanthellae in bleached regions of a colony exhibit reduced photosynthetic activity as long as one year after a bleaching event and that in situ fluorescence techniques such as FRRF are an effective means of studying coral responses and recovery from natural or anthropogenic stress in a non-destructive manner.     

The Effect of Bleaching on the Terpene Chemistry of Plexaurella fusifera: Evidence that Zooxanthellae Are Not Responsible for Sesquiterpene Production

The close association between marine invertebrates, zooxanthellae, and numerous bacteria gives rise to the question of the identity of the actual producer of secondary metabolites. In fall of 2005, a widespread bleaching event occurred throughout the Caribbean Sea in which some colonies of the gorgonian coral Plexaurella fusifera bleached. This study investigated whether zooxanthellae play a key role in the biosynthesis of secondary metabolite terpenes from P. fusifera. The extent of bleaching was examined by chlorophyll A analysis and also by zooxanthellae isolation and cell counting. The bleached and unbleached colonies were found to contain similar concentrations of eremophilene as the major terpene, and both exhibited similar biosynthetic capability as evaluated by the transformation of [C(1)-(3)H]-farnesyl diphosphate to the sesquiterpenes. Differences in bacterial communities between the bleached and unbleached colonies were analyzed using molecular techniques, and preliminary indications are that unbleached and bleached corals are dominated by low G + C firmicutes and gammaproteobacteria, respectively. It therefore appears that terpene biosynthesis can proceed independently of the zooxanthellae in P. fusifera, suggesting that the coral or a bacterium is the biosynthetic source.     

Bleaching increases likelihood of disease on Acropora palmata (Lamarck) in Hawksnest Bay, St John, US Virgin Islands

Anomalously high water temperatures may enhance the likelihood of coral disease outbreaks by increasing the abundance or virulence of pathogens, or by increasing host susceptibility. This study tested the compromised-host hypothesis, and documented the relationship between disease and temperature, through monthly monitoring of Acropora palmata colonies from May 2004 to December 2006, in Hawksnest Bay, St John, US Virgin Islands (USVI). Disease prevalence and the rate of change in prevalence showed a positive linear relationship with water temperature and rate of change in water temperature, respectively, but only in 2005 during prolonged periods of elevated temperature. Both bleached and unbleached colonies showed a positive relationship between disease prevalence and temperature in 2005, but the average area of disease-associated mortality increased only for bleached corals, indicating host susceptibility, rather than temperature per se, influenced disease severity on A. palmata.     

Coral microbiome diversity reflects mass coral bleaching susceptibility during the 2016 El Niño heat wave

Repeat marine heat wave‐induced mass coral bleaching has decimated reefs in Seychelles for 35 years, but how coral‐associated microbial diversity (microalgal endosymbionts of the family Symbiodiniaceae and bacterial communities) potentially underpins broad‐scale bleaching dynamics remains unknown. We assessed microbiome composition during the 2016 heat wave peak at two contrasting reef sites (clear vs. turbid) in Seychelles, for key coral species considered bleaching sensitive (Acropora muricata, Acropora gemmifera) or tolerant (Porites lutea, Coelastrea aspera). For all species and sites, we sampled bleached versus unbleached colonies to examine how microbiomes align with heat stress susceptibility. Over 30% of all corals bleached in 2016, half of which were from Acropora sp. and Pocillopora sp. mass bleaching that largely transitioned to mortality by 2017. Symbiodiniaceae ITS2‐sequencing revealed that the two Acropora sp. and P. lutea generally associated with C3z/C3 and C15 types, respectively, whereas C. aspera exhibited a plastic association with multiple D types and two C3z types. 16S rRNA gene sequencing revealed that bacterial communities were coral host‐specific, largely through differences in the most abundant families, Hahellaceae (comprising Endozoicomonas), Rhodospirillaceae, and Rhodobacteraceae. Both Acropora sp. exhibited lower bacterial diversity, species richness, and community evenness compared to more bleaching‐resistant P. lutea and C. aspera. Different bleaching susceptibility among coral species was thus consistent with distinct microbiome community profiles. These profiles were conserved across bleached and unbleached colonies of all coral species. As this pattern could also reflect a parallel response of the microbiome to environmental changes, the detailed functional associations will need to be determined in future studies. Further understanding such microbiome‐environmental interactions is likely critical to target more effective management within oceanically isolated reefs of Seychelles.     

Cellular processes of bleaching in the Mediterranean coral Oculina patagonica

Annual bleaching of Oculina patagonica on the Israeli Mediterranean coastline has been reported since 1993, although the cellular mechanisms underlying the bleaching have not yet been investigated. This survey examined 48 coral colonies of O. patagonica (bleached and unbleached) from various sites along the Israeli coast. Histopathological investigations of bleached lesions revealed a loss of endosymbionts, and an apparent in situ degradation of the endosymbionts. In situ end labelling of bleaching lesions did not provide evidence of apoptotic cell death. Electron microscopy of bleaching lesions also demonstrated an apparent in situ degradation and no evidence of apoptotic cell death of the host.     

Impacts of bleaching on the soft coral Lobophytum compactum. I. Fecundity, fertilization and offspring viability

We document long-term effects of a simulated bleaching event on the reproductive output and offspring viability of the soft coral Lobophytum compactum. Corals were subjected to temperature and solar radiation treatments to produce both moderately (48–60%) and heavily (90–95%) bleached colonies. Although bleached colonies recovered their zooxanthellae within 10 to 18 weeks, impacts on reproductive output were significant for at least two annual spawning seasons. In the first year, both polyp fecundity and mean oocyte diameter were reduced and inversely correlated with the degree of bleaching, with complete failure of fertilization in the group of heavily bleached colonies. For moderately bleached soft corals, survival and growth of sexual offspring did not differ significantly from those of unbleached colonies. Although no further reductions in zooxanthellae densities in experimental soft corals were recorded throughout the subsequent second year, egg size and fecundity of the heavily bleached soft corals were still significantly reduced 20 months later. Severe bleaching clearly has long-term sub-lethal impacts, reducing overall reproductive output for at least two spawning seasons. Accepted: 1 June 2000     

Coscinaraea marshae corals that have survived prolonged bleaching exhibit signs of increased heterotrophic feeding

Colonies of Coscinaraea marshae corals from Rottnest Island, Western Australia have survived for more than 11 months in various bleached states following a severe heating event in the austral summer of 2011. These colonies are situated in a high-latitude, mesophotic environment, which has made their long-term survival of particular interest as such environments typically suffer from minimal thermal pressures. We have investigated corals that remain unbleached, moderately bleached, or severely bleached to better understand potential survival mechanisms utilised in response to thermal stress. Specifically, Symbiodinium (algal symbiont) density and genotype, chlorophyll-a concentrations, and δ¹³C and δ¹⁵N levels were compared between colonies in the three bleaching categories. Severely bleached colonies housed significantly fewer Symbiodinium cells (p < 0.05) and significantly reduced chlorophyll-a concentrations (p < 0.05), compared with unbleached colonies. Novel Symbiodinium clade associations were observed for this coral in both severely and moderately bleached colonies, with clade C and a mixed clade population detected. In unbleached colonies, only clade B was observed. Levels of δ¹⁵N indicate that severely bleached colonies are utilising heterotrophic feeding mechanisms to aid survival whilst bleached. Collectively, these results suggest that these C. marshae colonies can survive with low symbiont and chlorophyll densities, in response to prolonged thermal stress and extended bleaching, and increase heterotrophic feeding levels sufficiently to meet energy demands, thus enabling some colonies to survive and recover over long time frames. This is significant as it suggests that corals in mesophotic and high-latitude environments may possess considerable plasticity and an ability to tolerate and adapt to large environmental fluctuations, thereby improving their chances of survival as climate change impacts coral ecosystems worldwide.     

Differences in microbial communities between healthy and bleached coral Acropora solitaryensis from Xisha Islands,South China Sea

Serious bleaching events have been observed in Acropora solitaryensis, one of the main species of reef-building coral in the area of the Xisha islands in the South China Sea, during 2008–2011. The microbial communities of healthy and bleached coral samples were compared to explore the difference in the bacterial composition using 16S rRNA gene sequencing and denaturing gradient gel electrophoresis (DGGE). Analysis of 16S rRNA gene sequences showed that the diversity of bacteria from the corals was different between the healthy and the bleached. Albeit both the healthy coral and bleached coral displayed similar dominant bacterial species (α- and γ-proteobacteria), the ratio of Vibrio spp. increased sharply in the latter. As the bleaching developed, the diversity of the microbial community was dramatically decreased and the dominant species were replaced by γ-proteobacteria where Vibrio spp. and Escherichia spp. overwhelmed other genera. Similar results were gained by the DGGE technique though the abundance was lower. Furthermore, in the bleached tissues, Vibrio coralliilyticus was identified using both methods. These results indicated that pathogenic Vibrio spp. might be one of the factors causing A. solitaryensis bleaching in the coral reef of the Xisha islands.     

Photoacclimatory and photoprotective responses to cold versus heat stress in high latitude reef corals

Corals at the world's southernmost coral reef of Lord Howe Island (LHI) experience large temperature and light fluctuations and need to deal with periods of cold temperature (<18°C), but few studies have investigated how corals are able to cope with these conditions. Our study characterized the response of key photophysiological parameters, as well as photoacclimatory and photoprotective pigments (chlorophylls, xanthophylls, and β‐carotene), to short‐term (5‐d) cold stress (~15°C; 7°C below control) in three LHI coral species hosting distinct Symbiodinium ITS2 types, and compared the coral–symbiont response to that under elevated temperature (~29°C; 7°C above control). Under cold stress, Stylophora sp. hosting Symbiodinium C118 showed the strongest effects with regard to losses of photochemical performance and symbionts. Pocillopora damicornis hosting Symbiodinium C100/C118 showed less severe bleaching responses to reduced temperature than to elevated temperature, while Porites heronensis hosting Symbiodinium C111* withstood both reduced and elevated temperature. Under cold stress, photoprotection in the form of xanthophyll de‐epoxidation increased in unbleached P. heronensis (by 178%) and bleached Stylophora sp. (by 225%), while under heat stress this parameter increased in unbleached P. heronensis (by 182%) and in bleached P. damicornis (by 286%). The xanthophyll pool size was stable in all species at all temperatures. Our comparative study demonstrates high variability in the bleaching vulnerability of these coral species to low and high thermal extremes and shows that this variability is not solely determined by the ability to activate xanthophyll de‐epoxidation.     

Structural Differences in Unbleached and Mildly‐Bleached Synthetic Tyrosine‐Derived Melanins Identified by Scanning Probe Microscopies

Using scanning tunneling microscopy (STM), we have imaged two types of mildly‐bleached, synthetic tyrosine‐derived melanins for comparison with the unbleached melanin from which they were prepared. These mildly‐bleached melanins were generated by mild oxidation of the unbleached melanin, using either basic hydrogen peroxide or air/light. The unbleached melanin, and two mildly‐bleached melanins, were independently deposited from very dilute tetrahydrofuran (THF) solutions onto highly oriented pyrolytic graphite (HOPG) substrate for STM imaging. Lateral dimensions (23 Å, average of two directions) of structures from each of the three samples showed no differences. However, structures from both mildly‐bleached melanins showed similar dramatic decreases (from ～15 Å to ～5 Å) in their STM‐measured apparent heights, compared with structures from the unbleached melanin sample. These STM observations are compatible with structural models for unbleached and mildly‐bleached melanins, incorporating a three‐dimensional structure for unbleached melanin composed of multi‐layered, Π–Π‐stacked, carboxylic and amino variants of polyaromatic polymeric sheets. The STM‐observed decrease in apparent heights after mild oxidation, which we associate with a change in stack height, has been confirmed by experiments using tapping mode atomic force microscopy (TM‐AFM) for the unbleached and mildly‐hydrogen‐peroxide‐bleached melanins (from ～14 Å to ～6 Å). In these TM‐AFM experiments, the melanins were deposited directly onto magnesium cation‐treated glass substrates in contact with methanolic solutions of each of the melanins. We interpret our mild‐bleaching results as an oxidative conversion of the multi‐layered, stacked sheets of mainly carboxylic and amino variants of polyquinhydrone‐like moieties, to largely de‐stacked, mildly‐bleached melanin sheets. These oxidized and, hence, electron‐deficient sheets should not readily form multi‐layered, Π–Π interacting stacks, but instead appear to be either single‐layer polyquinone sheets or, at most, double‐layer polyquinhydrone sheets. The effects of such de‐stacking on in vivo melanin photoprotection, and structural similarities between melanin derived from natural sources and the synthetic melanin samples used in this work are discussed.     

The effects of prolonged “bleaching” on the tissue biomass and reproduction of the reef coral Montastrea annularis

Colonies of Montastrea annularis from Carysfort Reef, Florida, that remained bleached seven months after the 1987 Caribbean bleaching event were studied to determine the long term effects of bleaching on coral physiology. Two types of bleached colonies were found: colonies with low numbers of zooxanthellae with normal pigment content, and a colony with high densities of lowpigment zooxanthellae. In both types, the zooxanthellae had an abnormal distribution within polyp tissues: highest densities were observed in basal endoderm and in mesenteries where zooxanthellae are not normally found. Bleached corals had 30% less tissue carbon and 44% less tissue nitrogen biomass per skeletal surface area, but the same tissue C:N ratio as other colonies that either did not bleach (normal) or that bleached and regained their zooxanthellae (recovered). Bleached corals were not able to complete gametogenesis during the reproductive season following the bleaching, while recovered corals were able to follow a normal gametogenic cycle. It appears that bleached corals were able to survive the prolonged period without nutritional contribution from their zooxanthellae by consuming their own structural materials for maintenance, but then, did not have the resources necessary for reproduction. The recovered corals, on the other hand, must have regained their zooxanthellae soon after the bleaching event since neither their tissue biomass nor their ability to reproduce were impaired.     

Influence of habitat degradation on fish replenishment

Temperature-induced coral bleaching is a major threat to the biodiversity of coral reef ecosystems. While reductions in species diversity and abundance of fish communities have been documented following coral bleaching, the mechanisms that underlie these changes are poorly understood. The present study examined the impacts of coral bleaching on the early life-history processes of coral reef fishes. Daily monitoring of fish settlement patterns found that ten times as many fish settled to healthy coral than sub-lethally bleached coral. Species diversity of settling fishes was least on bleached coral and greatest on dead coral, with healthy coral having intermediate levels of diversity. Laboratory experiments using light-trap caught juveniles showed that different damselfish species chose among healthy, bleached and dead coral habitats using different combinations of visual and olfactory cues. The live coral specialist, Pomacentrus moluccensis, preferred live coral and avoided bleached and dead coral, using mostly visual cues to inform their habitat choice. The habitat generalist, Pomacentrus amboinensis, also preferred live coral and avoided bleached and dead coral but selected these habitats using both visual and olfactory cues. Trials with another habitat generalist, Dischistodus sp., suggested that vision played a significant role. A 20 days field experiment that manipulated densities of P. moluccensis on healthy and bleached coral heads found an influence of fish density on juvenile weight and growth, but no significant influence of habitat quality. These results suggests that coral bleaching will affect settlement patterns and species distributions by influencing the visual and olfactory cues that reef fish larvae use to make settlement choices. Furthermore, increased fish density within the remaining healthy coral habitats could play an important role in influencing population dynamics.     

IMPORTANCE OF MACRO‐ VERSUS MICROSTRUCTURE IN MODULATING LIGHT LEVELS INSIDE CORAL COLONIES1

Adjusting the light exposure and capture of their symbiotic photosynthetic dinoflagellates (genus Symbiodinium Freud.) is central to the success of reef‐building corals (order Scleractinia) across high spatio‐temporal variation in the light environment of coral reefs. We tested the hypothesis that optical properties of tissues in some coral species can provide light management at the tissue scale comparable to light modulation by colony architecture in other species. We compared within‐tissue scalar irradiance in two coral species from the same light habitat but with contrasting colony growth forms: branching Stylophora pistillata and massive Lobophyllia corymbosa. Scalar irradiance at the level of the symbionts (2 mm into the coral tissues) were <10% of ambient irradiance and nearly identical for the two species, despite substantially different light environments at the tissue surface. In S. pistillata, light attenuation (90% relative to ambient) was observed predominantly at the colony level as a result of branch‐to‐branch self‐shading, while in L. corymbosa, near‐complete light attenuation (97% relative to ambient) was occurring due to tissue optical properties. The latter could be explained partly by differences in photosynthetic pigment content in the symbiont cells and pigmentation in the coral host tissue. Our results demonstrate that different strategies of light modulation at colony, polyp, and cellular levels by contrasting morphologies are equally effective in achieving favorable irradiances at the level of coral photosymbionts.     

Development of microsatellite markers as a molecular tool for conservation studies of the Mediterranean reef builder coral Cladocora caespitosa (Anthozoa, Scleractinia)

Cladocora caespitosa is a reef-building zooxanthellate scleractinian coral in the Mediterranean Sea. Mortality events have recurrently affected this species during the last decade. Thus, knowledge of its genetic structure, population diversity, and connectivity is needed to accomplish suitable conservation plans. In order to obtain a better understanding of the population genetics of this species, 13 highly variable microsatellites markers were developed from a naturally bleached colony. The developed primers failed to amplify zooxanthella DNA, isolated from C. caespitosa, verifying that these markers were of the coral and not algal symbiont origin. The degree of polymorphism of these loci was tested on tissue samples from 28 colonies. The allele number for each loci ranged from 2 to 13 (mean N(a) = 5.4), with an average observed heterozygosity of 0.42 (H(e) = 0.43) and all loci were in Hardy-Weinberg equilibrium. These new markers should be useful in future conservation genetic studies and will help to improve the resolution of the individual identification within this coral species. Primers were also tested in Oculina patagonica, with successful amplifications of several loci.     

Localized bleaching in Hawaii causes tissue loss and a reduction in the number of gametes in <Emphasis Type="Italic">Porites compressa</Emphasis>

Localized bleaching (a discrete white area on the coral) was observed in one of the main framework-building corals in Hawaii, Porites compressa. This study aimed to determine the degree of virulence of the lesion. We investigated the whole-colony effects by following disease progression through time and examining the effect of localized bleaching on coral fecundity. After two months, 35 of 42 (83.3%) individually tagged colonies affected by localized bleaching showed tissue loss and partial colony mortality. Histological slides of healthy P. compressa and samples from colonies showing signs of localized bleaching were compared showing that affected colonies had a significant reduction (almost 50%) in gamete development, egg numbers, and egg size in the affected tissue. The observed localized bleaching results in both partial colony mortality and a reduced number of gametes and was termed Porites Bleaching with Tissue Loss (PBTL).     

Effects of disturbance on coral communities: bleaching in Moorea,French Polynesia

This study examines patterns of susceptibility and short-term recovery of corals from bleaching. A mass coral bleaching event began in March, 1991 on reefs in Moorea, French Polynesia and affected corals on the shallow barrier reef and to >20 m depth on the outer forereef slope. There were significant differences in the effect of the bleaching among common coral genera, with Acropora, Montastrea, Montipora, and Pocillopora more affected than Porites, Pavona, leptastrea or Millepora. Individual colonies of the common species of Acropora and Pocillopora were marked and their fate assessed on a subsequent survey in August, 1991 to determine rates of recovery and mortality. Ninety-six percent of Acropora spp. showed some degree of bleaching compared to 76% of Pocillopora spp. From March to August mortality of bleached colonies of Pocillopora was 17%, 38% recovered completely, and many suffered some partial mortality of the tissue. In contrast, 63% of the Acropora spp. died, and about 10% recovered completely. Generally, those colonies with less than 50% of the colony area affected by the bleaching recovered at a higher rate than did those with more severe bleaching. Changes in community composition four months after the event began included a significant decrease only in crustose algae and an increase in cover of filamentous algae, much of which occupied plate-like and branching corals that had died in the bleaching event. Total coral cover and cover of susceptible coral genera had declined, but not significantly, after the event.     

Calcification in bleached and unbleached Montastraea faveolata: evaluating the role of oxygen and glycerol

All reef-building corals are symbiotic with dinoflagellates of the genus Symbiodinium, which influences many aspects of the host’s physiology including calcification. Coral calcification is a biologically controlled process performed by the host that takes place several membranes away from the site of photosynthesis performed by the symbiont. Although it is well established that light accelerates CaCO₃ deposition in reef-building corals (commonly referred to as light-enhanced calcification), the complete physiological mechanism behind the process is not fully understood. To better comprehend the coral calcification process, a series of laboratory experiments were conducted in the major Caribbean reef-building species Montastraea faveolata, to evaluate the effect of glycerol addition and/or the super-saturation of oxygen in the seawater. These manipulations were performed in bleached and unbleached corals, to separate the effect of photosynthesis from calcification. The results suggest that under normal physiological conditions, a 42% increase in seawater oxygen concentration promotes a twofold increase in dark-calcification rates relative to controls. On the other hand, the results obtained using bleached corals suggest that glycerol is required, as a metabolic fuel, in addition to an oxygenic environment in a symbiosis that has been disrupted. Also, respiration rates in symbiotic corals that were pre-incubated in light conditions showed a kinetic limitation, whereas corals that were pre-incubated in darkness were oxygen limited, clearly emphasizing the role of oxygen in this regard. These findings indicate that calcification in symbiotic corals is not strictly a “light-enhanced” or “dark-repressed” process, but rather, the products of photosynthesis have a critical role in calcification, which should be viewed as a “photosynthesis-driven” process. The results presented here are discussed in the context of the current knowledge of the coral calcification process.