Experimental assessment of the feeding effort of three scleractinian coral species during a thermal stress: Effect on the rates of photosynthesis

This study aimed at investigating changes in feeding rates of three scleractinian coral species (Stylophora pistillata, Turbinaria reniformis and Galaxea fascicularis) between control (26 °C) and short-term stress conditions (31 °C), and to assess the effect of feeding on the photosynthetic efficiency of the corals. Feeding rates varied according to the feeding effort of the corals, itself depending on the environmental conditions. Indeed, S. pistillata significantly decreased its feeding rates at 31 °C, while rates of T. reniformis and G. fascicularis were increased between 26 and 31 °C. Independently of the feeding rates, food supply helped in preventing damage to the photosynthetic apparatus of the zooxanthellae. Indeed, starved corals from the three species showed significant decrease in both the electron transport rates and in the photosynthetic rates, following a loss in the amount of chlorophyll and experiencing photoinhibition of the photosystem II. However, no bleaching was observed in heated fed corals, with no decrease in their photosynthetic efficiency or performance.     

Photo-acclimation of the hermatypic coral Stylophora pistillata while subjected to either starvation or food provisioning

This study investigated the photo-acclimation capacity of the coral Stylophora pistillata (Esper). Outer branches of coral colonies, taken from 2 m, were subjected to 90, 20, or 3% of incident surface photosynthetic active radiation (PAR(0)), or kept in total darkness. The corals were maintained either in filtered seawater (i.e., under starvation), or in seawater that had daily additions of zooplankton (rotifers). The experiments were maintained for 31 days. Zooxanthellae population densities and chlorophyll concentrations increased in S. pistillata fragments subjected to 20 and 3% PAR(0). The zooxanthellae densities decreased after 6 days in corals kept in total darkness, although chlorophyll concentrations remained higher. Corals that were fed and subjected to 90% PAR(0) showed lower degrading zooxanthellae frequencies, higher photosynthetic and respiration rates, and higher chlorophyll concentrations than corals in the same light regime under starvation. Complete acclimation to dim (20% PAR(0)) and low (3% PAR(0)) light was only apparent for corals fed with zooplankton. Changes in zooxanthellae population densities occurred through differential rates of zooxanthellae division and degradation.     

The effects of feeding frequency and symbiosis with zooxanthellae on the biochemical composition of Astrangia Danae Milne Edwards & Haime 1849

The coral Astrangia danae Milne Edwards & Haime 1849 occurs naturally with and without symbiotic algae and thus may have two sources of nourishment: (1) particles captured by the coral polyps, and (2) photosynthetic products translocated from their zooxanthellae. Symbiotic colonies may have both sources, and nonsymbiotic ones certainly have only the former. The relative importance of these two food sources was studied in the laboratory by examining the tissues of corals fed with frozen brine shrimp. Stock corals were fed once per week. Two to three weeks prior to each experiment, selected corals were placed on one of three feeding schedules: starved (S), fed once per week (1/wk), and fed three times per week (3/wk). The coral tissues were analyzed for protein, lipid, carbohydrate, and zooxanthellae content. Increased feeding frequency (1/wk → 3/wk) resulted in an increased tissue biomass and lipid to protein (L/P) ratio; starvation (1/wk → S) caused a decrease in these parameters. Symbiosis with zooxanthellae had an effect similar to increased feeding frequency in that the S and 1/wk symbiotic corals had a higher L/P ratio than comparable nonsymbiotic ones. There were no significant differences in L/P ratios between the 3/wk symbiotic and nonsymbiotic corals. Freshly collected colonies had a tissue composition most similar to the laboratory animals fed 3/wk. This result is consistent with the hypothesis that ingestion of solid food is the major nutritional source for A. danae in Narragansett Bay, Rhode Island, but our experiments suggest that the algae can have an important effect on tissue L/P ratios during times of food scarcity.     

Bioaccumulation of zinc in the scleractinian coral Stylophora pistillata

The uptake kinetics of zinc (Zn), an essential nutrient for both photosynthesis and calcification, in the tissue of S. pistillata showed that the transport of Zn is composed of a linear component (diffusion) at high concentrations and an active carrier-mediated component at low concentrations. The carrier affinity (K _m=28 pmol l⁻¹) was very low, indicating a good adaptation of the corals to low levels of Zn in seawater. Zn accumulation in the skeleton was linear; its level was dependent on the length of the incubation as well as on the external concentration of dissolved Zn. There was also a light-stimulation of Zn uptake, suggesting that zooxanthellae, through photosynthesis, are involved in this process. An enrichment of the incubation medium with 10 nM Zn significantly increased the photosynthetic efficiency of S. pistillata. This result suggests that corals living in oligotrophic waters might be limited in essential metals, such as zinc.     

Differential susceptibility to oxidative stress of two scleractinian corals: antioxidant functioning of mycosporine-glycine

This study examined the importance of mycosporine-glycine (Myc-Gly) as a functional antioxidant in the thermal-stress susceptibility of two scleractinian corals, Platygyra ryukyuensis and Stylophora pistillata. Photochemical efficiency of PSII (F_v/F_m), activity of antioxidant enzymes, superoxide dismutase (SOD) and catalase (CAT), and composition and abundance of mycosporine-like amino acids (MAAs) in the coral tissue and in symbiotic zooxanthellae were analyzed during 12-h exposure to high temperature (33 °C). After 6- and 12-h exposures at 33 °C, S. pistillata showed a significantly more pronounced decline in F_v/F_m compared to P. ryukyuensis. A 6-h exposure at 33 °C induced a significant increase in the activities of SOD and CAT in both host and zooxanthellae components of S. pistillata while in P. ryukyuensis a significant increase was observed only in the CAT activity of zooxanthellae. After 12-h exposure, the SOD activity of P. ryukyuensis was unaffected in the coral tissue but slightly increased in zooxanthellae, whereas the CAT activity in the coral tissue showed a 2.5-fold increase. The total activity of antioxidant enzymes was significantly higher in S. pistillata than in P. ryukyuensis, suggesting that P. ryukyuensis is less sensitive to oxidative stress than S. pistillata. This differential susceptibility of the corals is consistent with a 20-fold higher initial concentration of Myc-Gly in P. ryukyuensis compared to S. pistillata. In the coral tissue and zooxanthellae of both species investigated, the first 6 h of exposure to thermal stress induced a pronounced reduction in the abundance of Myc-Gly but not in other MAAs. When exposure was prolonged to 12 h, the Myc-Gly pool continued to decrease in P. ryukyuensis and was completely depleted in S. pistillata. The delay in the onset of oxidative stress in P. ryukyuensis and the dramatic increase in the activities of the antioxidant enzymes in S. pistillata, which contains low concentrations of Myc-Gly suggest that Myc-Gly provides rapid protection against oxidative stress before the antioxidant enzymes are induced. These findings strongly suggest that Myc-Gly is functioning as a biological antioxidant in the coral tissue and zooxanthellae and demonstrate its importance in the survival of reef-building corals under thermal stress.     

Impacts of Sediments on Coral Energetics: Partitioning the Effects of Turbidity and Settling Particles

Sediment loads have long been known to be deleterious to corals, but the effects of turbidity and settling particles have not previously been partitioned. This study provides a novel approach using inert silicon carbide powder to partition and quantify the mechanical effects of sediment settling versus reduced light under a chronically high sedimentary regime on two turbid water corals commonly found in Singapore (Galaxea fascicularis and Goniopora somaliensis). Coral fragments were evenly distributed among three treatments: an open control (30% ambient PAR), a shaded control (15% ambient PAR) and sediment treatment (15% ambient PAR; 26.4 mg cm⁻² day⁻¹). The rate of photosynthesis and respiration, and the dark-adapted quantum yield were measured once a week for four weeks. By week four, the photosynthesis to respiration ratio (P/R ratio) and the photosynthetic yield (F_v/F_m) had fallen by 14% and 3–17% respectively in the shaded control, contrasting with corals exposed to sediments whose P/R ratio and yield had declined by 21% and 18–34% respectively. The differences in rates between the shaded control and the sediment treatment were attributed to the mechanical effects of sediment deposition. The physiological response to sediment stress differed between species with G. fascicularis experiencing a greater decline in the net photosynthetic yield (13%) than G. somaliensis (9.5%), but a smaller increase in the respiration rates (G. fascicularis = 9.9%, G. somaliensis = 14.2%). These different physiological responses were attributed, in part, to coral morphology and highlighted key physiological processes that drive species distribution along high to low turbidity and depositional gradients.     

Feeding sustains photosynthetic quantum yield of a scleractinian coral during thermal stress

Thermal resistance of the coral-zooxanthellae symbiosis has been associated with chronic photoinhibition, increased antioxidant activity and protein repair involving high demands of nitrogen and energy. While the relative importance of heterotrophy as a source of nutrients and energy for cnidarian hosts, and as a means of nitrogen acquisition for their zooxanthellae, is well documented, the effect of feeding on the thermal sensitivity of the symbiotic association has been so far overlooked. Here we examine the effect of zooplankton feeding versus starvation on the bleaching susceptibility and photosynthetic activity of photosystem II (PSII) of zooxanthellae in the scleractinian coral Stylophora pistillata in response to thermal stress (daily temperature rises of 2-3 degrees C) over 10 days, employing pulse-amplitude-modulated chlorophyll fluorometry. Fed and starved corals displayed a decrease in daily maximum potential quantum yield (F (v)/F (m)) of PSII, effective quantum yield (F/F (m)') and relative electron transport rates over the course of 10 days. However after 10 days of exposure to elevated temperature, F (v)/F (m) of fed corals was still 50-70% higher than F (v)/F (m) of starved corals. Starved corals showed strong signs of chronic photoinhibition, which was reflected in a significant decline in nocturnal recovery rates of PSII relative to fed corals. This was paralleled by the progressive inability to dissipate excess excitation energy via non-photochemical quenching (NPQ). After 10 days, NPQ of starved corals had decreased by about 80% relative to fed corals. Feeding treatment had no significant effect on chlorophyll a and c (2) concentrations and zooxanthellae densities, but the mitotic indices were significantly lower in starved than in fed corals. Collectively the results indicate that exogenous food may reduce the photophysiological damage of zooxanthellae that typically leads to bleaching and could therefore play an important role in mediating the thermal resistance of some corals.     

Urea uptake by the scleractinian coral Stylophora pistillata

Urea can be one of the major sources of nitrogen for phytoplankton, but little is known about its importance for corals. Experiments were therefore designed to assess the uptake rates of urea by the scleractinian coral Stylophora pistillata; ¹⁵N-urea was used to follow the incorporation of nitrogen into the zooxanthellae and animal tissue. The uptake kinetics of urea in the tissue of S. pistillata showed that there is a concentration-dependent uptake of urea. The transport of urea was composed of a linear component (diffusion) at concentrations higher than 6 μmol N-urea l^− 1 and an active carrier-mediated component, at lower concentrations. The value of the carrier affinity (K_m = 1.05 μmol urea l^− 1) indicates a good adaptation of the corals to low levels of urea in seawater. At the in situ concentration of ca. 0.2 μmol N-urea l^− 1, the uptake rate was equal to 0.1 nmol N h^− 1 cm^− 2. Urea uptake was at least four times higher in the animal than in the algal fraction, and five times higher when corals were incubated in the light than in the dark. These results could be explained by the involvement of urea in the calcification process, which is also enhanced by light. Comparison of urea uptake rates with nitrate or ammonium uptake rates for the same S. pistillata species, at in situ concentrations, showed that urea is preferred to nitrate and may therefore be an important source of nitrogen for scleractinian corals.     

Effect of zooplankton availability on the rates of photosynthesis, and tissue and skeletal growth in the scleractinian coral Stylophora pistillata

This work investigated the effect of light and feeding on tissue composition as well as on rates of photosynthesis and calcification in the zooxanthellae (zoox) scleractinian coral, Stylophora pistillata. Microcolonies were maintained at three different light levels (80, 200, 300 μmol m⁻² s⁻¹) and subjected to two feeding regimes (starved and fed) over 9 weeks. Corals were fed both natural plankton and Artemia salina nauplii four times a weeks and samplings were made after 2, 5, and 9 weeks. Results confirmed that feeding enhances coral growth rate and increases both the dark and light calcification rates. These rates were 50-75% higher in fed corals (FC; 60±20 and 200±40 nmol Ca²⁺ cm⁻² h⁻¹ for dark and light calcification, respectively) compared to control corals (CC; 30±9 and 124±23 nmol Ca²⁺ cm⁻² h⁻¹). The dark calcification rates, however, were four times lower than the rates of light calcification (independent of trophic status). After 5 weeks, chlorophyll a (chl-a) concentrations were four to seven times higher in fed corals (7-21 μg cm⁻²) than in control corals (2-5 μg cm⁻²). The amount of protein was also significantly higher in fed corals (2.11-2.50 mg cm⁻²) than in control corals (1.08-1.52 mg cm⁻²). Rates of photosynthesis in fed corals were 2-10 times higher (1.24±0.75 μmol O₂ h⁻¹ cm⁻²) than those measured in control corals (0.20±0.08 μmol O₂ h⁻¹ cm⁻²).     

Symbiosis-specific changes in dimethylsulphoniopropionate concentrations in Stylophora pistillata along a depth gradient

Scleractinian corals are prolific producers of dimethylsulphoniopropionate (DMSP), but ecophysiological mechanisms influencing cellular concentrations are uncertain. While DMSP is often proposed to function as an antioxidant, interactions between specific host–symbiont genotype associations, plasticity in DMSP concentrations and environmental conditions that can either exert or alleviate oxidative stress are unclear. We used long-term (6 months) reciprocal transplantation of Stylophora pistillata hosting two distinct symbiont phylotypes along a depth gradient, clades A (<20 m) and C (>20 m), to assess the effect of change in depth (light intensity) on DMSP concentrations in relation to symbiont genotype and photoacclimation in corals between 3 and 50 m in the Gulf of Aqaba. Bathymetric distribution of total DMSP (DMSPt) per cell varied significantly while particulate DMSP (DMSPp) appeared to be unaffected by depth. Highest DMSPt concentrations in control corals occurred at 20 m. While 3-m transplants showed a significant increase in DMSPt concentration at 20 m and became affiliated with an additional genotype (C72), 50-m transplants largely persisted with their original genotype and exhibited no significant changes in DMSPt concentrations. DMSPt concentrations in transplants at both 3 and 50 m, on the other hand, increased significantly while all corals maintained their original symbiont genotypes. Photoacclimation differed significantly with transplantation direction relative to the controls. Symbionts in 3-m transplants at 20 m exhibited no changes in chlorophyll a (chl a) concentration, cell density or cell diameter while symbiont densities decreased and chl a concentrations increased significantly at 50 m. In contrast, symbiont densities in 50-m transplants remained unaffected across depths while symbiont diameters decreased. Chl a concentrations decreased at 20 m and increased at 3 m. Our results indicate that DMSPt concentrations following changes in depth are not only a function of symbiont genotype but result from different acclimation abilities of both symbiotic partners.

     

Size variation of planulae and its effect on the lifetime of planulae in three pocilloporid corals

Dispersal of propagules plays an important role in the distribution of corals. Pocillopora damicornis, Seriatopora hystrix, and Stylophora pistillata are all brooders and release planulae having symbiotic zooxanthellae. Planulae showed a great size variation, especially at peaks of planulation, and we found negative correlations between zooxanthella density and planula size in S. hystrix and S. pistillata. Studies of the larval life of planulae under both light and dark conditions have revealed that larger planulae have a longer lifetime. When planulae of the same size were compared, it was found that they lived longer under light conditions than under dark conditions. These findings suggest that planulae utilize energy from photosynthetic products of zooxanthellae and that these corals enjoy long-distance dispersal by producing larger planulae with greater dispersal potential. It is conceivable that variation in the dispersal potential of planulae is a means of adaptation by which planulae can increase their chances of finding a suitable habitat.     

Effects of host nutritional status and seasonality on the nitrogen status of zooxanthellae in the temperate coral Plesiastrea versipora (Lamarck)

The nitrogen status of endosymbiotic dinoflagellates (zooxanthellae) in the temperate coral Plesiastrea versipora (Lamarck) was determined by measuring the extent to which ammonium (40 μM NH₄⁺) enhanced the rate of zooxanthellar dark carbon fixation above that seen in filtered seawater (FSW) alone; the enhancement ratio was expressed as [dark NH₄⁺ rate/dark FSW rate]. V_D′/V_L, a further index of nitrogen status, was also calculated where V_D′ = [dark NH₄⁺ rate − dark FSW rate] and V_L = rate of carbon fixation in the light. When corals were starved for 2-8 weeks, zooxanthellar nitrogen deficiency became apparent at ≥ 4 weeks, with NH₄⁺/FSW and V_D′/V_L averaging up to 2.08 and 0.0061, respectively. A decrease in light-saturated photosynthesis per zooxanthella also occurred, with the photosynthetic rate after 4-6 weeks being just 81% of that seen prior to starvation. In comparison, when corals were fed 5 times per week for 8 weeks the addition of ammonium had little effect, indicating nitrogen sufficiency; NH₄⁺/FSW and V_D′/V_L were 1.03 and 0.0003, respectively. Photosynthetic rates of zooxanthellae from well-fed corals were up to 1.7 times greater than those of zooxanthellae from starved corals. The nitrogen status of zooxanthellae from corals in the field exhibited seasonal differences. Autumn samples were nitrogen sufficient, with NH₄⁺/FSW = 1.003 and V_D′/V_L = 0.0001. In contrast, a small degree of nitrogen deficiency was seen in winter and spring, when NH₄⁺/FSW averaged 1.075 and 1.249, and V_D′/V_L averaged 0.0013 and 0.0014, respectively. The greatest degree of nitrogen deficiency was observed in summer, when NH₄⁺/FSW averaged 1.318 and V_D′/V_L averaged 0.0036. Given the clear links between food supply and nitrogen status seen under experimental conditions, and the likelihood that the zooxanthellae are also able to take up nutrients directly from the seawater, the fluctuations in nitrogen status may reflect temporal fluctuations in seawater nutrient concentrations and plankton abundance. The nutrient status of these temperate zooxanthellae in the field is in contrast to the marked nitrogen deficiency seen in zooxanthellae from nutrient-poor coral reef waters, and raises the possibility that temperate zooxanthellae can store nitrogen for use when exogenous nutrients and food are less readily available. This, in turn, may contribute to the considerable stability of temperate zooxanthellar populations under highly variable environmental conditions.     

Effect of natural zooplankton feeding on the tissue and skeletal growth of the scleractinian coral<Emphasis Type="Italic"> Stylophora pistillata</Emphasis>

Laboratory experiments were designed to estimate the ingestion rates of the scleractinian coral Stylophora pistillata under varying prey concentrations and feeding regimes and to assess the effect of feeding on the tissue and skeletal growth. Six sets of corals were incubated under two light (80 and 300 µmol photons m^–2 s^–1) and three feeding levels (none, fed twice, and fed six times per week) using freshly collected zooplankton. Results showed that the number of prey ingested was proportional to prey density, and no saturation of feeding capability was reached. Capture rates varied between 0.5 and 8 prey items 200 polyp^–1 h^–1. Corals starved for several days ingested more plankton than did fed corals. Fed colonies exhibited significantly higher levels of protein, chlorophyll a, and chlorophyll c₂ per unit surface area than starved colonies. Feeding had a strong effect on tissue growth, increasing it by two to eight times. Calcification rates were also 30% higher in fed than in starved corals. Even moderate levels of feeding enhanced both tissue and skeletal growth, although the processes involved in this enhancement remain to be determined.     

Physiological and Biochemical Performances of Menthol-Induced Aposymbiotic Corals

The unique mutualism between corals and their photosynthetic zooxanthellae (Symbiodinium spp.) is the driving force behind functional assemblages of coral reefs. However, the respective roles of hosts and Symbiodinium in this endosymbiotic association, particularly in response to environmental challenges (e.g., high sea surface temperatures), remain unsettled. One of the key obstacles is to produce and maintain aposymbiotic coral hosts for experimental purposes. In this study, a simple and gentle protocol to generate aposymbiotic coral hosts (Isopora palifera and Stylophora pistillata) was developed using repeated incubation in menthol/artificial seawater (ASW) medium under light and in ASW in darkness, which depleted more than 99% of Symbiodinium from the host within 4∼8 days. As indicated by the respiration rate, energy metabolism (by malate dehydrogenase activity), and nitrogen metabolism (by glutamate dehydrogenase activity and profiles of free amino acids), the physiological and biochemical performances of the menthol-induced aposymbiotic corals were comparable to their symbiotic counterparts without nutrient supplementation (e.g., for Stylophora) or with a nutrient supplement containing glycerol, vitamins, and a host mimic of free amino acid mixture (e.g., for Isopora). Differences in biochemical responses to menthol-induced bleaching between Stylophora and Isopora were attributed to the former digesting Symbiodinium rather than expelling the algae live as found in the latter species. Our studies showed that menthol could successfully bleach corals and provided aposymbiotic corals for further exploration of coral-alga symbioses.     

Inhibition of coral and algal photosynthesis by ca-antagonist phenothiazine drugs

The effects of various calcium ion antagonists and ion transport inhibitors on photosynthetic O₂ evolution of corals, isolated zooxanthellae, sea anemone tentacles, and Chlorococcum oleofaciens were measured. Only the phenothiazine drugs were effective at inhibiting photosynthesis. Trifluoperazine, a calcium ion antagonist drug, inhibited at low concentrations, with 10⁻⁴ molar and 8 × 10⁻⁶ molar completely abolishing photosynthesis in the intact corals and isolated zooxanthellae, respectively. Net photosynthetic O₂ evolution of C. oleofaciens was eliminated by concentrations of trifluoperazine as low as 2.8 × 10⁻⁵ molar.     

Inhibition of Calvin–Benson cycle suppresses the repair of photosystem II in Symbiodinium: implications for coral bleaching

Inhibition of Calvin–Benson cycle (CBC) activity by thermal stress has been hypothesized to cause photoinhibition of photosystem II (PSII) in zooxanthellae of reef-building corals and consequently lead to bleaching. This study tests whether the interruption of CBC by glycolaldehyde (GA) leads to photoinhibition and subsequent coral bleaching in Stylophora pistillata. When S. pistillata was incubated with GA, the O₂ evolution rate declined in a dose-dependent manner and the extent of photoinhibition, reflected by a decreased maximum quantum yield of PSII (F _v/F _m), was enhanced. The effect of GA on photoinhibition was similar to that of chloramphenicol (CAP), an inhibitor of protein synthesis in chloroplasts. When S. pistillata was incubated in weak light following a high-light-induced photoinhibitory treatment, the recovery of PSII from photoinhibition was suppressed in a similar manner to both GA- and CAP-treated samples. After incubation in moderate light at 26°C, S. pistillata showed a bleaching response only in presence of GA. These results suggest that coral bleaching-like responses are caused by interruption of the CBC activity in S. pistillata and are associated with accelerated photoinhibition through suppression of the protein synthesis-dependent repair of PSII but not to an increase in photodamage to PSII.     

Nutritional status and metabolism of the coral Stylophora subseriata along a eutrophication gradient in Spermonde Archipelago (Indonesia)

Coral responses to degrading water quality are highly variable between species and depend on their trophic plasticity, acclimatization potential, and stress resistance. To assess the nutritional status and metabolism of the common scleractinian coral, Stylophora subseriata, in situ experiments were carried along a eutrophication gradient in Spermonde Archipelago, Indonesia. Coral fragments were incubated in light and dark chambers to measure photosynthesis, respiration, and calcification in a number of shallow reefs along the gradient. Chlorophyll a (chl a), protein content, maximum quantum yield (F _v/F _m), and effective quantum yield (Φ PS II) were measured on the zooxanthellae, in addition to host tissue protein content and biomass. Photosynthetic rates were 2.5-fold higher near-shore than mid-shelf due to higher areal zooxanthellae and chl a concentrations and a higher photochemical efficiency (Φ PS II). A 2- and 3-fold increase in areal host tissue protein and biomass was found, indicating a higher nutritional supply in coastal waters. Dark respiration, however, showed no corresponding changes. There was a weak correlation between calcification and photosynthesis (Pearson r = 0.386) and a lack of metabolic stress, as indicated by constant respiration and F _v/F _m and the “clean” and healthy appearance of the colonies in spite of high turbidity in near-shore waters. The latter suggests that part of the energetic gains through increased auto- and heterotrophy were spent on metabolic expenditures, e.g., mucus production. While coastal pollution is always deleterious to the reef ecosystem as a whole, our results show that the effect on corals may not always be negative. Thus, S. subseriata may be one of the few examples of corals actually profiting from land-based sources of pollution.     

Response of two species of Indo-Pacific corals, Porites cylindrica and Stylophora pistillata, to short-term thermal stress: The host does matter in determining the tolerance of corals to bleaching

The role of both host and dinoflagellate symbionts was investigated in the response of reef-building corals to thermal stress in the light. Replicate coral nubbins of Stylophora pistillata and Porites cylindrica from the GBR were exposed to either 28 °C (control) or 32 °C for 5 days before being returned to an ambient reef temperature (28 °C). S. pistillata was found to contain either Symbiodinium genotype C1 or C8a, while P. cylindrica had type C15 based on ITS genotyping. Analysis of the quantum yield of photosystem (PS) II fluorescence of the symbionts in P. cylindrica showed that light-induced excitation pressure on the C15 Symbiodinium was significantly less, and the steady state quantum yield of PSII fluorescence at noon (ΔF/Fm′) greater, than that measured in C1/C8a Symbiodinium sp. from S. pistillata. Immunoblots of the PS II D1 protein were significantly lower in Symbiodinium from S. pistillata compared to those in P. cylindrica after exposure to thermal stress. The biochemical markers, heat-stress protein (HSP) 70 and superoxide dismutase (SOD), were significantly greater in P. cylindrica before the experiment, and both species of coral increased their biosynthesis of HSP 70 and SOD when exposed to thermal stress. Concentrations of MAAs, glycerol, and lipids were not significantly affected by thermal stress in these experiments, but DNA damage was greater in heat-stressed S. pistillata compared to P. cylindrica. There was minimal coral mucus, which accounts for up to half of the total energy budget of a coral and provides the first layer of defense for invading microbes, produced by S. pistillata after heat stress compared to P. cylindrica. It is concluded that P. cylindrica contains a heat resistant C15 Symbiodinium and critical host proteins are present at higher concentrations than observed for S. pistillata, the combination of which provides greater protection from bleaching conditions of high temperature in the light.     

Regulation and control of intracellular algae (= zooxanthellae) in hard corals

To examine algal (= zooxanthellae) regulation and control, and the factors determining algal densities in hard corals, the zooxanthellae mitotic index and release rates were regularly determined in branch tips from a colony of a staghorn coral, Acropora formosa, recovering from a coral ''bleaching'' event (the stress-related dissociation of the coral–algal symbiosis). Mathematical models based upon density-dependent decreases in the algal division frequency and increases in algal release rates during the post-bleaching recovery period accurately predict the observed recovery period (ca. 20 weeks). The models suggest that (i) the colony recovered its algal population from the division of the remaining zooxanthellae, and (ii) the continual loss of zooxanthellae significantly slowed the recovery of the coral. Possible reasons for the ''paradoxical'' loss of healthy zooxanthellae from the bleached coral are discussed in terms of endodermal processes occurring in the recovering coral and the redistribution of newly formed zooxanthellae to aposymbiotic host cells. At a steady-state algal density of 2.1 x 10⁶ zooxanthellae cm^-2 at the end of the recovery period, the zooxanthellae would have to form a double layer of cells in the coral tissues, consistent with microscopic observations. Neighbouring colonies of A. formosa with inherently higher algal densities possess proportionately smaller zooxanthellae. Results suggest that space availability and the size of the algal symbionts determines the algal densities in the coral colonies. The large increases in the algal densities reported in corals exposed to elevated nutrient concentrations (i.e between a two- and five-fold increase in the algal standing stock) are not consistent with this theory. We suggest that increases of this magnitude are a product of the experimental conditions: reasons for this statement are discussed. We propose that the stability of the coral–algal symbiosis under non-stress conditions, and the constancy of zooxanthellae densities in corals reported across growth form, depth and geographic range, are related to space availability limiting algal densities. However, at these densities, zooxanthellae have attributes consistent with nutrient limitation.     

Recovery of the coral Montastrea annularis in the Florida Keys after the 1987 Caribbean “bleaching event”

Many reef-building corals and other cnidarians lost photosynthetic pigments and symbiotic algae (zooxanthellae) during the coral bleaching event in the Caribbean in 1987. The Florida Reef Tract included some of the first documented cases, with widespread bleaching of the massive coral Montastrea annularis beginning in late August. Phototransects at Carysfort Reef showed discoloration of >90% of colonies of this species in March 1988 compared to 0% in July 1986; however no mortality was observed between 1986 and 1988. Samples of corals collected in February and June 1988 had zooxanthellae densities ranging from 0.1 in the most lightly colored corals, to 1.6x10⁶ cells/cm² in the darker corals. Minimum densities increased to 0.5x10⁶ cells/cm² by August 1989. Chlorophyll-a content of zooxanthellae and zooxanthellar mitotic indices were significantly higher in corals with lower densities of zooxanthellae, suggesting that zooxanthellar at low densities may be more nutrientsufficient than those in unbleached corals. Ash-free dry weight of coral tissue was positively correlated with zooxanthellae density at all sample times and was significantly lower in June 1988 compared to August 1989. Proteins and lipids per cm² were significantly higher in August 1989 than in February or June, 1988. Although recovery of zooxanthellae density and coral pigmentation to normal levels may occur in less than one year, regrowth of tissue biomass and energy stores lost during the period of low symbiont densities may take significantly longer.