Threatened Caribbean Coral Is Able to Mitigate the Adverse Effects of Ocean Acidification on Calcification by Increasing Feeding Rate

Global climate change threatens coral growth and reef ecosystem health via ocean warming and ocean acidification (OA). Whereas the negative impacts of these stressors are increasingly well-documented, studies identifying pathways to resilience are still poorly understood. Heterotrophy has been shown to help corals experiencing decreases in growth due to either thermal or OA stress; however, the mechanism by which it mitigates these decreases remains unclear. This study tested the ability of coral heterotrophy to mitigate reductions in growth due to climate change stress in the critically endangered Caribbean coral Acropora cervicornis via changes in feeding rate and lipid content. Corals were either fed or unfed and exposed to elevated temperature (30°C), enriched pCO₂ (800 ppm), or both (30°C/800 ppm) as compared to a control (26°C/390 ppm) for 8 weeks. Feeding rate and lipid content both increased in corals experiencing OA vs. present-day conditions, and were significantly correlated. Fed corals were able to maintain ambient growth rates at both elevated temperature and elevated CO₂, while unfed corals experienced significant decreases in growth with respect to fed conspecifics. Our results show for the first time that a threatened coral species can buffer OA-reduced calcification by increasing feeding rates and lipid content.     

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).     

The importance of zooplankton to the daily metabolic carbon requirements of healthy and bleached corals at two depths

Bleached and non-bleached fragments of three species of Hawaiian corals were exposed to enhanced and ambient concentrations of zooplankton at 1 and 6 m depth to determine the contribution of zooplankton to the coral's daily carbon budget. The size and taxonomic grouping were recorded for every zooplankton captured and the relative input of zooplankton of different size classes was determined. The contribution of heterotrophy to animal respiration (CHAR) was calculated using an improved method that included the proportionate contribution of zooplankton from all size classes. Results show that the proportionate effects of species, depth and bleaching treatments on coral feeding rates were not significantly different between ambient and enhanced zooplankton concentrations. Corals captured the same size and assemblage of zooplankton under all evaluated conditions, and preferentially captured plankters smaller than 400 µm. Feeding rates of Porites lobata increased with depth regardless of bleaching status. Feeding rates of Porites compressa increased with depth in non-bleached corals, but not in bleached corals. Within depth, feeding rates of bleached Montipora capitata increased, P. compressa decreased and P. lobata remained unchanged relative to non-bleached fragments. Therefore, the feeding response of corals to the same disturbance may vary considerably. Calculated CHAR values show that heterotrophic carbon from zooplankton plays a much larger role in the daily carbon budget of corals than previously estimated, accounting for 46% of some coral species' daily metabolic carbon requirements when healthy and 147% when bleached. Thus, heterotrophically acquired carbon made an important contribution to the daily carbon budget of corals under all experimental conditions. These results suggest that the relative importance of autotrophic and heterotrophic carbon to a coral's energetic needs is mediated by a coral's bleaching status and environment, and should be considered on a continuum, from 100% photoautotrophy to 100% heterotrophy.     

Calcification by juvenile corals under heterotrophy and elevated CO2

Ocean acidification (OA) threatens the existence of coral reefs by slowing the rate of calcium carbonate (CaCO₃) production of framework-building corals thus reducing the amount of CaCO₃ the reef can produce to counteract natural dissolution. Some evidence exists to suggest that elevated levels of dissolved inorganic nutrients can reduce the impact of OA on coral calcification. Here, we investigated the potential for enhanced energetic status of juvenile corals, achieved via heterotrophic feeding, to modulate the negative impact of OA on calcification. Larvae of the common Atlantic golf ball coral, Favia fragum, were collected and reared for 3 weeks under ambient (421 μatm) or significantly elevated (1,311 μatm) CO₂ conditions. The metamorphosed, zooxanthellate spat were either fed brine shrimp (i.e., received nutrition from photosynthesis plus heterotrophy) or not fed (i.e., primarily autotrophic). Regardless of CO₂ condition, the skeletons of fed corals exhibited accelerated development of septal cycles and were larger than those of unfed corals. At each CO₂ level, fed corals accreted more CaCO₃ than unfed corals, and fed corals reared under 1,311 μatm CO₂ accreted as much CaCO₃ as unfed corals reared under ambient CO₂. However, feeding did not alter the sensitivity of calcification to increased CO₂; ? calcification/?Ω was comparable for fed and unfed corals. Our results suggest that calcification rates of nutritionally replete juvenile corals will decline as OA intensifies over the course of this century. Critically, however, such corals could maintain higher rates of skeletal growth and CaCO₃ production under OA than those in nutritionally limited environments.     

The biology and physiology of alga-invertebrate symbioses. III. In situ measurements of photosynthesis and calcification in some hermatypic corals

In situ measurements of the rates of photosynthesis and calcification in three species of hermatypic corals were made at Eilat, in the Gulf of Aqaba, Red Sea. Experiments were made at 5, 20 and 35 m depth under unusually poor conditions of submarine illumination for the region, and at the relatively low water temperature (21°C) for coral growth which prevails there all year. Estimates of photosynthetic rates by both the ¹⁴C and oxygen methods indicated that the ¹⁴C method does measure gross photosynthesis in these organisms even at, and below, light compensation points. Substantial rates of carbon fixation in Acropora and Millepora show that, even under bad conditions, these organisms could survive autotrophically to at least 10 m depth, as also could the massive coral Goniastrea although this had much lower photosynthetic rates under the same conditions, compensated for by a much lower respiratory rate than the other two corals.Calcification rates were variable but showed a considerable increase in light as compared with the dark in all three species, and the rates did not decrease with depth as much as might have been anticipated from the reduction in photosynthesis and ambient light energy. Photosynthetic and calcification rates were similar to those reported for similar organisms both in the Caribbean and on the Great Barrier Reef.     

Effects of coral bleaching on the feeding response of two species of coral-feeding fish

Coral bleaching is an increasingly prominent threat to coral reef ecosystems, not only to corals, but also to the many organisms that rely on coral for food and shelter. Coral-feeding fishes are negatively affected by coral loss caused by extensive bleaching, but it is unknown how feeding behaviour of most corallivorous fishes changes in response to coral bleaching. In this study, coral bleaching was experimentally induced in situ to examine the feeding response of two obligate corallivorous fish, Labrichthys unilineatus (Labridae) and Chaetodon baronessa (Chaetodontidae). Feeding rates were monitored before, during, and immediately after experimental bleaching of prey corals. L. unilineatus significantly increased its feeding on impacted corals during bleaching, but showed a steady decline in feeding once corals were fully bleached. Feeding response of L. unilineatus appears to parallel the expected stress-induced mucous production by bleaching colonies. In contrast, C. baronessa preferentially fed from healthy colonies over bleached colonies, although bleached colonies were consumed for five days following manipulation. Feeding by corallivorous fishes can play an important role in determining coral condition and mortality of corals following stress induced bleaching.     

Carbon translocation from symbiont to host depends on irradiance and food availability in the tropical coral Stylophora pistillata

Reef-building corals live in symbiosis with dinoflagellates that translocate a large proportion of their photosynthetically fixed carbon compounds to their coral host for its own metabolism. The carbon budget and translocation rate, however, vary depending on environmental conditions, coral host species, and symbiont clade. To quantify variability in carbon translocation in response to environmental conditions, this study assessed the effect of two different irradiance levels (120 and 250 μmol photons m^?2 s^?1) and feeding regimes (fed with Artemia salina nauplii and unfed) on the carbon budget of the tropical coral Stylophora pistillata. For this purpose, H¹³CO₃ ^?-enriched seawater was used to trace the conversion of photosynthetic carbon into symbiont and coral biomass and excrete particulate organic carbon. Results showed that carbon translocation (ca. 78 %) and utilization were similar under both irradiance levels for unfed colonies. In contrast, carbon utilization by fed colonies was dependent on the growth irradiance. Under low irradiance, heterotrophy was accompanied by lower carbon translocation (71 %), higher host and symbiont biomass, and higher calcification rates. Under high irradiance, heterotrophy was accompanied by higher rates of photosynthesis, respiration, and carbon translocation (90 %) as well as higher host biomass. Hence, levels of resource sharing within coral–dinoflagellate symbioses depend critically on environmental conditions.     

Effects of temperature, light and heterotrophy on the growth rate and budding of the temperate coral Cladocora caespitosa

Recent investigations have shown the temperate scleractinian coral Cladocora caespitosa to be a new potential climate archive for the Mediterranean Sea. Whilst earlier studies have demonstrated a seasonal variation in growth rates, they were unable to distinguish which environmental parameter (light, temperature, or food) was influencing growth. In this study, the effect of these three factors on the coral physiology and calcification rate was characterized to aid the correct interpretation of skeletal trace element variations. Two temperatures (13 and 23°C), irradiances (50 and 120 μmol m⁻² s⁻¹), and feeding regimes (unfed and fed with nauplii of Artemia salina) were tested under controlled laboratory conditions on the growth, zooxanthellae density, chlorophyll (chl) content, and asexual reproduction (budding) of C. caespitosa during a 7-week factorial experiment. Unlike irradiance, which had no effect, high temperature and food supply increased the growth rates of C. caespitosa. The effect of feeding was however higher for corals maintained at low temperature, suggesting that heterotrophy is especially important during the cold season, and that temperature is the predominant factor affecting the coral’s growth. At low temperature, fed samples had higher zooxanthellae density and chl content, possibly for maximizing photosynthetic efficiency. Sexual reproduction investment of C. caespitosa was higher during favourable conditions characterised by high temperatures and zooplankton availability.     

Effect of naturally changing zooplankton concentrations on feeding rates of two coral species in the Eastern Pacific

Zooplankton concentrations are known to vary by as much as an order of magnitude over a lunar cycle. Here, we conducted an experiment to determine the effect of ambient zooplankton concentrations over a lunar cycle on feeding rates of the corals Pavona gigantea (Verrill) (mounding coral, 3.0 mm diameter polyps) and Pocillopora damicornis (Linnaeus) (branching coral, 1.0 mm diameter polyps) in situ on a shallow reef at Isla Contadora, Gulf of Panamá (Pacific), Panamá. Coral fragments exposed to either enhanced or ambient zooplankton concentrations were allowed to feed for 1 h, collected, and their gut contents dissected. The number of zooplankton captured was counted, feeding rates calculated per cm², and the species composition of captured zooplankton assemblages determined. Although both species captured the same zooplankton assemblage, feeding rates were always significantly higher for P. gigantea than for P. damicornis. Under ambient flow and zooplankton concentrations, feeding rates were highly correlated with zooplankton concentration in the 200-400 μm size class. Under constantly enhanced zooplankton concentrations in the control fragments, feeding rates did not vary significantly over the lunar cycle. As such, coral feeding rates vary not as a result of lunar phase per se, but with changes in zooplankton abundance over the lunar cycle. Coral feeding rates are directly proportional to ambient zooplankton concentrations and may vary by as much as 50% over a lunar cycle, suggesting that corals must cope with major swings in sources of fixed carbon and nutrients over relatively short timescales.     

Temperature effects on calcification rate and skeletal deposition in the temperate coral, Plesiastrea versipora (Lamarck)

The geographic range of the coral, Plesiastrea versipora (Lamarck, 1816), extends into temperate waters outside the southern limit for hermatypic corals. In the present study, calcification in Plesiastrea collected from Port Phillip Bay, Victoria was examined over the coral's normal annual temperature range (10-21 °C), which is well below the normal optimum for coral calcification in tropical corals (25-28 °C). Calcification rate in Plesiastrea was considerably lower than in reef corals, but showed a similar pattern in temperature responses, with a trend towards higher rates at ∼18 °C. The light/dark calcification ratio was markedly lower than that in tropical corals. Autoradiography showed that calcification occurred primarily by deposition of calcium carbonate at the upper surfaces of the septo-costae. Scanning electron microscopy (SEM) showed that skeletal deposition in Plesiastrea had a temperature-dependent diel pattern. In the light, calcium carbonate was deposited as small spheroidal crystals and, at higher temperatures, small needle-shaped crystals. In the dark, calcium carbonate deposition appeared to be in the form of an amorphous sheet-like cementation. Compared with other scleractinian corals, calcification rate in Plesiastrea was relatively slow and showed different patterns of skeletal deposition.     

The rôle of zooxanthellae in the hermatypic coral Plesiastrea urvillei (Milne Edwards and Haime) From cold waters

The presence of zooxanthellae in tissues of the cold-temperate water coral Plesiastrea urvillei (Milne Edwards and Haime) has been confirmed histologically. Numbers of zooxanthellae per unit surface area and increases in submerged wet weight as a measure of calcification have been followed for 150 days under four different conditions: light-fed, light-starved, dark-fed, and dark-starved. No significant difference was found in density of zooxanthellae or calcification rates between light-fed and light-starved, and between dark-fed and dark-starved. After Day 48 the calcification rate in the dark dropped, however, by a factor of ≈4 to a constant lower rate and was correlated with a decrease in density of zooxanthellae. Zooxanthellae thus enhance calcification about 4 times during photosynthesis. Measurements of oxygen consumption and production indicated that even at the low light intensities experienced on a cloudy winter day by the coral in its natural environment more energy was fixed during photosynthesis than was required by the host. The retention of zooxanthellae and continued calcification in the dark for upwards of 48 days is considered to be an adaptation to the lower light levels experienced by P. urvillei compared with tropical corals.     

Coral photobiology: new light on old views

The relationship between reef-building corals and light-harvesting pigments of zooxanthellae (Symbiodinium sp.) has been acknowledged for decades. The photosynthetic activity of the algal endocellular symbionts may provide up to 90% of the energy needed for the coral holobiont. This relationship limits the bathymetric distribution of coral reefs to the upper 100 m of tropical shorelines. However, even corals growing under high light intensities have to supplement the photosynthates translocated from the algae by predation on nutrient-rich zooplankton. New information has revealed how the fate of carbon acquired through photosynthesis differs from that secured by predation, whose rates are controlled by light-induced tentacular extension. The Goreau paradigm of “light-enhanced calcification” is being reevaluated, based on evidence that blue light stimulates coral calcification independently from photosynthesis rates. Furthermore, under dim light, calcification rates were stoichiometrically uncoupled from photosynthesis. The rates of photosynthesis of the zooxanthellae exhibit a clear endogenous rhythmicity maintained by light patterns. This daily pattern is concomitant with a periodicity of all the antioxidant protective mechanisms that wax and wane to meet the concomitant fluctuation in oxygen evolution. The phases of the moon are involved in the triggering of coral reproduction and control the spectacular annual mass-spawning events taking place in several reefs. The intensity and directionality of the underwater light field affect the architecture of coral colonies, leading to an optimization of the exposure of the zooxanthellae to light. We present a summary of major gaps in our understanding of the relationship between light and corals as a roadmap for future research.     

The effect of heterotrophy on photosynthesis and tissue composition of two scleractinian corals under elevated temperature

Exogenous food can increase protein levels of coral host tissue, zooxanthellae densities, chlorophyll (chl) concentrations and rates of photosynthesis and is thought to play an important role in the resilience of bleached corals. There is however no information about the effect of heterotrophy on the bleaching susceptibility of corals under elevated temperature conditions. This study investigates potential interactions between food availability, basal metabolic functions (photosynthesis and respiration), energy status (lipid concentrations), total protein concentrations and the bleaching susceptibility (loss of chl and/or zooxanthellae) of the scleractinian corals Stylophora pistillata (Esper) and Galaxea fascicularis (Linnaeus) in response to elevated temperature (daily temperature rises of 3-4 °C) over 15 days. Feeding experiments were carried out in which the corals were either fed daily with zooplankton or starved. Compared to fed corals, starvation of both species resulted in a significant decrease in daily photosynthetic oxygen evolution over time. Gross (Pg) and net (Pn) photosynthetic production of starved corals of both species between 10:00-11:00 hrs had declined by ~50% at day 15 while there were no marked changes in Pg and Pn of fed corals. After 15 days, starved S. pistillata contained significantly lower zooxanthellae densities, lipid and protein concentrations than fed corals. Starved G. fascicularis also displayed a decrease in zooxantllae densities which was accompanied by a significant decline in algal chl concentrations. Contrary to S. pistillata, feeding treatment had no effect on the lipid concentrations of G. fascicularis. Total protein concentrations however were significantly lower in straved than in fed G. fascicularis. Furthermore, starvation resulted in a significant decrease in respiration of S. pistillata during the last four days of the experiment while treatment had no effect on the respiration rates of G. fascicularis. Overall the oxygen consumption of S. pistillata of both treatments was about 39-67% higher than the respiration of G. fascicularis indicating that low metabolic rates may have allowed starved G. fascicularis to conserve energy reserves over the course of the experiment. The combined results reveal a strong positive relationship between food availability, sustained photosynthetic activity and reduced loss in pigmentation of both species under elevated temperature 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.     

Heterotrophy in Tropical Scleractinian Corals

The dual character of corals, that they are both auto- and heterotrophs, was recognized early in the twentieth Century. It is generally accepted that the symbiotic association between corals and their endosymbiotic algae (called zooxanthellae) is fundamental to the development of coral reefs in oligotrophic tropical oceans because zooxanthellae transfer the major part of their photosynthates to the coral host (autotrophic nutrition). However, numerous studies have confirmed that many species of corals are also active heterotrophs, ingesting organisms ranging from bacteria to mesozooplankton. Heterotrophy accounts for between 0 and 66% of the fixed carbon incorporated into coral skeletons and can meet from 15 to 35% of daily metabolic requirements in healthy corals and up to 100% in bleached corals. Apart from this carbon input, feeding is likely to be important to most scleractinian corals, since nitrogen, phosphorus, and other nutrients that cannot be supplied from photosynthesis by the coral's symbiotic algae must come from zooplankton capture, particulate matter or dissolved compounds. A recent study showed that during bleaching events some coral species, by increasing their feeding rates, are able to maintain and restore energy reserves.
This review assesses the importance and effects of heterotrophy in tropical scleractinian corals. We first provide background information on the different food sources (from dissolved organic matter to meso- and macrozooplankton). We then consider the nutritional inputs of feeding. Finally, we review feeding effects on the different physiological parameters of corals (tissue composition, photosynthesis and skeletal growth).     

Dark calcification and the daily rhythm of calcification in the scleractinian coral, <Emphasis Type="Italic">Galaxea fascicularis</Emphasis>

The rate of calcification in the scleractinian coral Galaxea fascicularis was followed during the daytime using ⁴⁵Ca tracer. The coral began the day with a low calcification rate, which increased over time to a maximum in the afternoon. Since the experiments were carried out under a fixed light intensity, these results suggest that an intrinsic rhythm exists in the coral such that the calcification rate is regulated during the daytime. When corals were incubated for an extended period in the dark, the calcification rate was constant for the first 4 h of incubation and then declined, until after one day of dark incubation, calcification ceased, possibly as a result of the depletion of coral energy reserves. The addition of glucose and Artemia reduced the dark calcification rate for the short duration of the experiment, indicating an expenditure of oxygen in respiration. Artificial hypoxia reduced the rate of dark calcification to about 25% compared to aerated coral samples. It is suggested that G. fascicularis obtains its oxygen needs from the surrounding seawater during the nighttime, whereas during the day time the coral exports oxygen to the seawater.     

Live tissue imaging shows reef corals elevate pH under their calcifying tissue relative to seawater

The threat posed to coral reefs by changes in seawater pH and carbonate chemistry (ocean acidification) raises the need for a better mechanistic understanding of physiological processes linked to coral calcification. Current models of coral calcification argue that corals elevate extracellular pH under their calcifying tissue relative to seawater to promote skeleton formation, but pH measurements taken from the calcifying tissue of living, intact corals have not been achieved to date. We performed live tissue imaging of the reef coral Stylophora pistillata to determine extracellular pH under the calcifying tissue and intracellular pH in calicoblastic cells. We worked with actively calcifying corals under flowing seawater and show that extracellular pH (pHe) under the calicoblastic epithelium is elevated by ～0.5 and ～0.2 pH units relative to the surrounding seawater in light and dark conditions respectively. By contrast, the intracellular pH (pHi) of the calicoblastic epithelium remains stable in the light and dark. Estimates of aragonite saturation states derived from our data indicate the elevation in subcalicoblastic pHe favour calcification and may thus be a critical step in the calcification process. However, the observed close association of the calicoblastic epithelium with the underlying crystals suggests that the calicoblastic cells influence the growth of the coral skeleton by other processes in addition to pHe modification. The procedure used in the current study provides a novel, tangible approach for future investigations into these processes and the impact of environmental change on the cellular mechanisms underpinning coral calcification.     

Symbiodinium Photosynthesis in Caribbean Octocorals

Symbioses with the dinoflagellate Symbiodinium form the foundation of tropical coral reef communities. Symbiodinium photosynthesis fuels the growth of an array of marine invertebrates, including cnidarians such as scleractinian corals and octocorals (e.g., gorgonian and soft corals). Studies examining the symbioses between Caribbean gorgonian corals and Symbiodinium are sparse, even though gorgonian corals blanket the landscape of Caribbean coral reefs. The objective of this study was to compare photosynthetic characteristics of Symbiodinium in four common Caribbean gorgonian species: Pterogorgia anceps, Eunicea tourneforti, Pseudoplexaura porosa, and Pseudoplexaura wagenaari. Symbiodinium associated with these four species exhibited differences in Symbiodinium density, chlorophyll a per cell, light absorption by chlorophyll a, and rates of photosynthetic oxygen production. The two Pseudoplexaura species had higher Symbiodinium densities and chlorophyll a per Symbiodinium cell but lower chlorophyll a specific absorption compared to P. anceps and E. tourneforti. Consequently, P. porosa and P. wagenaari had the highest average photosynthetic rates per cm² but the lowest average photosynthetic rates per Symbiodinium cell or chlorophyll a. With the exception of Symbiodinium from E. tourneforti, isolated Symbiodinium did not photosynthesize at the same rate as Symbiodinium in hospite. Differences in Symbiodinium photosynthetic performance could not be attributed to Symbiodinium type. All P. anceps (n = 9) and P. wagenaari (n = 6) colonies, in addition to one E. tourneforti and three P. porosa colonies, associated with Symbiodinium type B1. The B1 Symbiodinium from these four gorgonian species did not cluster with lineages of B1 Symbiodinium from scleractinian corals. The remaining eight E. tourneforti colonies harbored Symbiodinium type B1L, while six P. porosa colonies harbored type B1i. Understanding the symbioses between gorgonian corals and Symbiodinium will aid in deciphering why gorgonian corals dominate many Caribbean reefs.     

An experimental analysis of the effects of light and zooplankton on coral zonation

Summary In Panamá, vertical zonation of coral species is well marked; branching corals (Pocillopora spp.) predominate in shallow (1–6 m) water while massive forms (Pavona spp.) occupy the deeper (6–10 m) areas of the reef. To test the hypothesis that this zonation represents differential resource utilization (i.e., niche partitioning of food resources), one year manipulative field experiments were conducted at two depths (1 m and 7 m below Mean Lower Sea Level) assessing the relative contribution of light and zooplankton to the nutrition of three Pacific corals: Pocillopora damicornis, Pavona clavus and Pavona gigantea. Also tested were the related hypotheses that (i) energy for the maintenance and growth of corals comes mainly from light, independent of zooplankton supply and (ii) Porter's model, in which coral morphology is a predictor of the phototrophic-heterotrophic capabilities of a particular species. That is, corals with a branching morphology and a small polyp diameter (and short tentacle length) should be primarily phototrophic while those species with larger polyps and/or a massive form should show a greater degree of heterotrophy.The comparison of caged versus control corals indicated that the branching coral, Pocillopora damicornis (polyp diameter — 1 mm) grew independent of zooplankton supply (>95) but was markedly affected by shading. Pavona clavus (massive form with intermediate size polyps — 2 mm) was also negatively affected by shading but this effect was minimized when zooplankton was present. Surprisingly, under ambient light conditions, the growth of this specics was independent of zooplankton (>95); perhaps indicating a facultative reliance on zooplankton especially in shallow water. The third species, Pavona gigan-tea (massive form with largest polyps — 3 mm) was highly dependent on both light and zooplankton and was unable to compensate for the effects of shading by zooplankton feeding. This high dependence on light was unexpected since large polyps, according to Porter's model, would predict a more heterotrophic existence.Under all treatments, in both shallow and deep water, Pocillopora was found to grow more rapidly than either massive species. Thus, this study indicates that zonation on eastern Pacific reefs cannot be explained by depth-related differences in nutritional requirements among the species present. These results do provide the first long-term, experimental field evidence that corals are largely phototrophic organisms. The validity of Porter's model attempting to correlate phototrophic-heterotrophic abilities to morphological characteristics is, at least, partially verified: species with larger tentacles do appear to utilize zooplankton to a greater extent than those species with smaller tentacles. However, the significance of corallum morphology in predicting the mode of nutrient acquisition is unclear. Rather, behavioral patterns (tentacle expansion-contraction cycles) and location on the reef may be more important factors to consider.     

Coral bleaching is linked to the capacity of the animal host to supply essential metals to the symbionts

Massive coral bleaching events result in extensive coral loss throughout the world. These events are mainly caused by seawater warming, but are exacerbated by the subsequent decrease in nutrient availability in surface waters. It has therefore been shown that nitrogen, phosphorus or iron limitation contribute to the underlying conditions by which thermal stress induces coral bleaching. Generally, information on the trophic ecology of trace elements (micronutrients) in corals, and on how they modulate the coral response to thermal stress is lacking. Here, we demonstrate for the first time that heterotrophic feeding (i.e. the capture of zooplankton prey by the coral host) and thermal stress induce significant changes in micro element concentrations and isotopic signatures of the scleractinian coral Stylophora pistillata. The results obtained first reveal that coral symbionts are the major sink for the heterotrophically acquired micronutrients and accumulate manganese, magnesium and iron from the food. These metals are involved in photosynthesis and antioxidant protection. In addition, we show that fed corals can maintain high micronutrient concentrations in the host tissue during thermal stress and do not bleach, whereas unfed corals experience a significant decrease in copper, zinc, boron, calcium and magnesium in the host tissue and bleach. In addition, the significant increase in δ⁶⁵Cu and δ⁶⁶Zn signature of symbionts and host tissue at high temperature suggests that these isotopic compositions are good proxy for stress in corals. Overall, present findings highlight a new way in which coral heterotrophy and micronutrient availability contribute to coral resistance to global warming and bleaching.