Cell death and degeneration in the symbiotic dinoflagellates of the coral Stylophora pistillata during bleaching

Rising sea temperatures are increasing the incidences of mass coral bleaching (the dissociation of the coral-algal symbiosis) and coral mortality. In this study, the effects of bleaching (induced by elevated light and temperature) on the condition of symbiotic dinoflagellates (Symbiodinium sp.) within the tissue of the hard coral Stylophora pistillata (Esper) were assessed using a suite of techniques. Bleaching of S. pistillata was accompanied by declines in the maximum potential quantum yield of photosynthesis (F-v/F-m, measured using pulse amplitude modulated [PAM] fluorometry), an increase in the number of Sytox-green-stained algae (indicating compromised algal membrane integrity and cell death), an increase in 2',7'-dichlorodihydrofluroscein diacetate (H(2)DCFDA)stained algae (indicating increased oxidative stress), as well as ultrastructural changes (vacuolisation, losses of chlorophyll, and an increase in accumulation bodies). Algae expelled from S. pistillata exhibited a complete disorganisation of cellular contents; expelled cells contained only amorphous material. In situ samples taken during a natural mass coral bleaching event on the Great Barrier Reef in February 2002 also revealed a high number of Sytox-labelled algae cells in symbio. Dinoflagellate degeneration during bleaching seems to be similar to the changes resulting from senescence-phase cell death in cultured algae. These data support a role for oxidative stress in the mechanism of coral bleaching and highlight the importance of algal degeneration during the bleaching of a reef coral.

Testing the 'photoinhibition' model of coral bleaching using chemical inhibitors

Explants of the hard coral Seriatopora hystrix were exposed to sublethal concentrations of the herbicide diuron DCMU (N'-(3,4-dichlorophenyl,-N,N-dimethylurea)) and the heavy metal copper. Pulse amplitude modulated (PAM) chlorophyll fluorescence techniques were used to assess the effects on the photosynthetic efficiency of the algal symbionts in the tissue (in Symbio), and chlorophyll fluorescence and counts of symbiotic algae (normalised to surface area) were used to assess the extent of coral bleaching. At 30 mug DCMU l(-1), there was a reduction in both the maximum effective quantum yield (DeltaF/F-m') and maximum potential quantum yield (F-v/F-m) of the algal symbionts in symbio. Corals subsequently lost their algal symbionts and discoloured (bleached), especially on their upper sunlight-exposed surfaces. At the same DCMU concentration but under low light (5% of growth irradiance), there was a marked reduction in DeltaF/F-m' but only a slight reduction in F-v/F-m and slight loss of algae. Loss of algal symbionts was also noted after a 7 d exposure to concentrations as low as 10 mug DCMU l(-1) under normal growth irradiance, and after 14 d exposure to 10 mug DCMU l(-1) under reduced irradiance. Collectively the results indicate that DCMU-induced bleaching is caused by a light-dependent photoinactivation of algal symbionts, and that bleaching occurs when F-v/F-n, (measured 2 h after sunset) is reduced to a value of less than or equal to 0.6. Elevated copper concentrations (60 mug Cu l(-1) for 10 h) also induced a rapid bleaching in S. hystrix but without affecting the quantum yield of the algae in symbio. Tests with isolated algae indicated that substantially higher concentrations (300 mug Cu l(-1) for 8 h) were needed to significantly reduce the quantum yield. Thus, copper-induced bleaching occurs without affecting the algal photosynthesis and may be related to effects on the host (animal). It is argued that warm-water bleaching of corals resembles both types of chemically induced bleaching, suggesting the need for an integrated model of coral bleaching involving the effect of temperature on both host (coral) and algal symbionts.

Endolithic algae photoacclimate to increased irradiance during coral bleaching

The photoacclimation of endolithic algae ( of the genus Ostreobium) inhabiting the skeleton of the Mediterranean coral Oculina patagonica during a bleaching event was examined. Pulse amplitude modulated (PAM) chlorophyll fluorescence techniques in situ were used to assess the photosynthetic efficiency of endolithic algae in the coral skeleton and the symbiotic dinoflagellates (zooxanthellae) in the coral tissue. Relative photosynthetic electron transport rates (ETRs) of the endolithic algae under bleached areas of the colony were significantly higher than those of endolithic algae from a healthy section of the colony and those of zooxanthellae isolated from the same section. Endolithic algae under healthy parts of the colony demonstrated an ETRmax of 16.5% that of zooxanthellae from tissue in the same section whereas endolithic algae under bleached sections showed ETRmax values that were 39% of those found for healthy zooxanthellae. The study demonstrates that endolithic algae undergo photoacclimation with increased irradiance reaching the skeleton. As PAM fluorometry has become a major tool for assessing levels of stress and bleaching in corals, the importance of considering the contribution of the endolithic algae to the overall chlorophyll fluorescence measured is highlighted.

Dynamics of a temperature-related coral disease outbreak

During the austral summer of 2001/2002, a coral epizootic occurred almost simultaneously with a bleaching event on the fringing reefs of Magnetic Island (Great Barrier Reef region), Australia. This resulted in a 3- to 4-fold increase in the mean percentage of partial mortality rate in a population of the hard coral Montipora aequituberculata. The putative disease state, ‘atramentous necrosis’, was observed on both bleached and normally-pigmented M. aequituberculata, and presented blackened lesions that spread within days across the colony surface and throughout the population. Diseased portions of the corals were only visible for 3 to 4 wk, with diseased tissues becoming covered in sediment and algae, which rapidly obscured evidence of the outbreak. Diseased colonies were again observed in the summer of 2002/2003 after being absent over the 2002 winter. Analysis of when diseased and bleached corals were first observed, and when and where the mortality occurred on individual colonies, indicated virtually all the mortality over the summer could be attributed to the disease and not to the bleaching. Fluorescence in situ hybridisation (FISH) techniques and cloning, and analysis of the 16S rRNA genes from diseased coral tissue, identified a mixed microbial assemblage in the diseased tissues particularly within the Alphaproteobacteria, Firmicutes and Bacteroidetes. While it is not possible in this study to distinguish between a disease-causing microbial community versus secondary invaders, the bacterial 16S rDNA sequences identified within the blackened lesions demonstrated high similarity to sequences from black band disease and white plague infected corals, suggesting either common aetiological agents or development of a bacterial community that is specific to degrading coral tissues. Temperature-induced coral disease outbreaks, with the potential for elevated levels of mortality, may represent an added problem for corals during the warmer summer months and an added dimension to predicted increases in water temperature from climate change.

Estimates of sex- and age-class-specific survival probabilities for a southern Great Barrier Reef green sea turtle population

Sex- and age-class-specific survival probabilities of a southern Great Barrier Reef green sea turtle population were estimated using a capture - mark - recapture (CMR) study and a Cormack - Jolly - Seber (CJS) modelling approach. The CMR history profiles for 954 individual turtles tagged over a 9-year period ( 1984 - 1992) were classified into three age classes ( adult, subadult, juvenile) based on somatic growth and reproductive traits. Reduced-parameter CJS models, accounting for constant survival and time-specific recapture, fitted best for all age classes. There were no significant sex-specific differences in either survival or recapture probabilities for any age class. Mean annual adult survival was estimated at 0.9482 (95% CI: 0.92 - 0.98) and was significantly higher than survival for either subadults or juveniles. Mean annual subadult survival was 0.8474 ( 95% CI: 0.79 - 0.91), which was not significantly different from mean annual juvenile survival estimated at 0.8804 ( 95% CI: 0.84 - 0.93). The time-specific adult recapture probabilities were a function of sampling effort but this was not the case for either juveniles or subadults. The sampling effort effect was accounted for explicitly in the estimation of adult survival and recapture probabilities. These are the first comprehensive sex- and age-class-specific survival and recapture probability estimates for a green sea turtle population derived from a long-term CMR program.

Effects of the herbicide diuron on the early life history stages of coral

The effects of the herbicide diuron on the early life history stages of broadcast spawning and brooding corals were examined in laboratory experiments. Fertilisation of Acropora millepora and Montipora aequituberculata oocytes were not inhibited at diuron concentrations of up to 1000 mu gl(-1). Metamorphosis of symbiont-free A. millepora larvae was only significantly inhibited at 300 mu gl(-1) diuron. Pocillopora damicornis larvae, which contain symbiotic dinoflagellates, were able to undergo metamorphosis after 24h exposure to diuron at 1000 mu gl(-1). Two-week old P. damicornis recruits on the other hand were as susceptible to diuron as adult colonies, with expulsion of symbiotic dinoflagellates (bleaching) evident at 10 mu gl(-1) diuron after 96 h exposure. Reversible metamorphosis was observed at high diuron concentrations, with fully bleached polyps escaping from their skeletons. Pulse amplitude modulation (PAM) chlorophyll fluorescence techniques demonstrated a reduction in photosynthetic efficiency (Delta F/F'(m)) in illuminated P. dami- cornis recruits after a 2h exposure to 1 mu gl(-1) diuron. The dark-adapted quantum yields (F-v/F-m also declined, indicating chronic photoinhibition and damage to photosystem H. Crown Copyright (c) 2004 Published by Elsevier Ltd. All rights reserved.

Establishing representative no-take areas in the Great Barrier Reef : Large scale implementation of theory on marine protected areas

The Great Barrier Reef Marine Park, an area almost the size , of Japan, has a new network of no-take areas that significantly improves the protection of biodiversity. The new marine park zoning implements, in a quantitative manner, many of the theoretical design principles discussed in the literature. For example, the new network of no-take areas has at least 20% protection per bioregion, minimum levels of protection for all known habitats and special or unique features, and minimum sizes for no-take areas of at least 10 or 20 kat across at the smallest diameter Overall, more than 33% of the Great Barrier Reef Marine Park is now in no-take areas (previously 4.5%). The steps taken leading to this outcome were to clarify to the interested public why the existing level of protection wets inadequate; detail the conservation objectives of establishing new no-take areas; work with relevant and independent experts to define, and contribute to, the best scientific process to deliver on the objectives; describe the biodiversity (e.g., map bioregions); define operational principles needed to achieve the objectives; invite community input on all of The above; gather and layer the data gathered in round-table discussions; report the degree of achievement of principles for various options of no-take areas; and determine how to address negative impacts. Some of the key success factors in this case have global relevance and include focusing initial communication on the problem to be addressed; applying the precautionary principle; using independent experts; facilitating input to decision making; conducting extensive and participatory consultation; having an existing marine park that encompassed much of the ecosystem; having legislative power under federal law; developing high-level support; ensuring agency Priority and ownership; and being able to address the issue of displaced fishers.

Near-bottom depletion of zooplankton over coral reefs: III: vertical gradient of predation pressure

Many diurnal planktivorous fish in coral reefs efficiently consume zooplankton drifting in the overlying water column. Our survey, carried out at two coral reefs in the Red Sea, showed that most of the diurnal planktivorous fish foraged near the bottom, close to the shelters from piscivores. The planktivorous fish were order of magnitude more abundant near (

Allozyme electrophoresis still represents a powerful technique in the management of coral reefs

Understanding genetic variability and gene flow between populations of scleractinian corals separated by one to several hundred kilometers is crucially important as we head into a century of climate change in which an understanding of the connectivity of populations is a critically important question in management. Genetic methods that directly use molecular variance in the DNA should offer greater precision in detecting differences among individuals and populations than the more traditional allozyme electrophoresis. However, this paper highlights the point that the limited number of DNA markers that have been identified for scleractinian coral genetic studies do not necessarily offer greater precision than that offered by allozymes. In fact, at present allozyme electrophoresis yields greater information than the eight different DNA markers used in this study. Given the relative ease of use of allozymes and the wealth of comparable data sets from numerous previously published studies, allozyme electrophoresis should not be dismissed for population structure and connectivity studies on coral reefs. While continued effort should be placed into searching for new DNA markers, until a more sensitive DNA marker becomes available for scleractinian corals, allozyme electrophoresis remains a powerful and relevant technique for understanding the connectivity of coral population studies.

Coral bleaching following wintry weather

Extensive coral bleaching Occurred intertidally in early August 2003 in the Capricorn Bunker group (Wistari Reef, Heron and One Tree Islands; Southern Great Barrier Reef). The affected intertidal coral had been exposed to unusually cold (minimum = 13.3degreesC; wet bulb temperature = 9degreesC) and dry winds (44% relative humidity) for 2 d during predawn low tides. Coral bleached in the upper 10 cm of their branches and had less than 0.2 x 10(6) cell cm(-2) as compared with over 2.5 x 10(6), Cell cm(-2) in nonbleached areas. Dark-adapted quantum yields did not differ between symbionts in bleached and nonbleached areas. Exposing symbionts to light, however, led to greater quenching of Photosystem 11 in symbionts in the bleached coral. Bleached areas of the affected colonies had died by September 2003, with areas that were essentially covered by more than 80% living coral decreasing to less than 10% visible living coral cover. By January 2004, coral began to recover, principally from areas of colonies that were not exposed during low tide (i.e., from below dead, upper regions). These data highlight the importance of understanding local weather patterns as well as the effects of longer term trends in global climate.

A modern soft-bottom, shallow-water crinoid fauna (Echinodermata) from the Great Barrier Reef, Australia

A recent preliminary survey revealed that 12 species of unstalked crinoids occur on a gentle sandy slope (12-18 m depth) at Lizard Island, Great Barrier Reef, Australia; five of which are also found on coral reefs. The other seven appear to constitute a unique assemblage restricted to unconsolidated substrates, where most cling to algae or hide beneath rubble or sponges. Members of this assemblage exhibit all of the basic feeding postures found among reef-dwelling species. However, Comatula rotalaria, which lacks anchoring cirri and bears uniquely differentiated short and long arms, exhibits a posture different from other living crinoids. Quantitative transects reveal apparent depth-related differences in species composition: C. rotalaria dominated the 12 transects in 12-13 m (84% of 82 specimens), while Comatella nigra, Comatula cf. purpurea, Amphimetra cf. tessellata and Zygometra microdiscus accounted for 96% of 54 specimens observed along 12 transects in 16-17 m.

Diel 'tuning' of coral metabolism: physiological responses to light cues

Hermatypic-zooxanthellate corals track the diel patterns of the main environmental parameters temperature, UV and visible light - by acclimation processes that include biochemical responses. The diel course of solar radiation is followed by photosynthesis rates and thereby elicits simultaneous changes in tissue oxygen tension due to the shift in photosynthesis/respiration balance. The recurrent patterns of sunlight are reflected in fluorescence yields, photosynthetic pigment content and activity of the two protective enzymes superoxide dismutase (SOD) and catalase (CAT), enzymes that are among the universal defenses against free radical damage in living tissue. All of these were investigated in three scleractinian corals: Favia favus, Plerogyra sinuosa and Goniopora lobata. The activity of SOD and CAT in the animal host followed the course of solar radiation, increased with the rates of photosynthetic oxygen production and was correlated with a decrease in the maximum quantum yield of photochemistry in Photosystem H (PSII) (Delta F'/F-m'). SOD and CAT activity in the symbiotic algae also exhibited a light intensity correlated pattern, albeit a less pronounced one. The observed rise of the free-radical-scavenger enzymes, with a time scale of minutes to several hours, is an important protective mechanism for the existence and remarkable success of the unique cnidarian-dinoflagellate associations, in which photosynthetic oxygen production takes place within animal cells. This represents a facet of the precarious act of balancing the photosynthetic production of oxygen by the algal symbionts with their destructive action on all living cells, especially those of the animal host.

Ecological persistence interrupted in Caribbean coral reefs [Letter]

The recent mass mortality of Caribbean reef corals dramatically altered reef community structure and begs the question of the past stability and persistence of coral assemblages before human disturbance began. We report within habitat stability in coral community composition in the Pleistocene fossil record of Barbados for at least 95 000 years despite marked variability in global sea level and climate. Results were consistent for surveys of both common and rare taxa. Comparison of Pleistocene and modern community structure shows that Recent human impacts have changed coral community structure in ways not observed in the preceding 220 000 years.

Endoparasite communities of five fish species (Labridae : Cheilininae) from Lizard Island: how important is the ecology and phylogeny of the hosts?

The parasite community of animals is generally influenced by host physiology, ecology, and phylogeny. Therefore, sympatric and phylogenetically related hosts with similar ecologies should have similar parasite communities. To test this hypothesis we surveyed the endoparasites of 5 closely related cheilinine fishes (Labridae) from the Great Barrier Reef. They were Cheilinus chlorounts, C. trilobatus, C. fasciatils, Epibulus insidiator and OxYcheilinus diagrainnia. VVe examined the relationship between parasitological variables (richness, abundance and diversity) and host characteristics (bodv weight, diet and phuylogeny). The 5 fishes had 31 parasite species with 9-18 parasite species per fish species. Cestode larvae (mostly Tetraphyllidea) were the most abundant and prevalent parasites followed by nematodes and digeneans. Parasites, body size and diet of hosts differed between fish species. In general, body weight, diet and host phylogeny each explained some of the variation in richness and composition of parasites among the fishes. The 2 most closely related species, Cheilinus chlorourus and C. trilobatus, had broadly similar parasites but the Other fish species differed significantly in all variables. However, there was no all -encompassing pattern. This may, be because different lineages of parasites may react differently to ecological variables. We also argue that adult parasites may respond principally to host diet. In contrast, larval parasite composition may respond both to host diet and predator-prey interactions because this is the path by which many, parasites complete their life-cycles. Finally, variation in parasite phylogeny and parasite life-cycles among hosts likely increase the complexity of the system making it difficult to find all-encompassing patterns between host characteristics and parasites, particularly when all the species in rich parasite communities are considered.

Increased mortality and photoinhibition in the symbiotic dinoflagellates of the Indo-Pacific coral Stylophora pistillata (Esper) after summer bleaching

Coral bleaching (the loss of symbiotic dinoflagellates from reef-building corals) is most frequently caused by high-light and temperature conditions. We exposed the explants of the hermatypic coral Stylophora pistillata to four combinations of light and temperature in late spring and also in late summer. During mid-summer, two NOAA bleaching warnings were issued for Heron Island reef (Southern Great Barrier Reef, Australia) when sea temperature exceeded the NOAA bleaching threshold, and a 'mild' (in terms of the whole coral community) bleaching event occurred, resulting in widespread S. pistillata bleaching and mortality. Symbiotic dinoflagellate biomass decreased by more than half from late spring to late summer (from 2.5x10(6) to 0.8x10(6) dinoflagellates cm(2) coral tissue), and those dinoflagellates that remained after summer became photoinhibited more readily (dark-adapted F (V) : F (M) decreased to (0.3 compared with 0.4 in spring), and died in greater numbers (up to 17% dinoflagellate mortality compared with 5% in the spring) when exposed to artificially elevated light and temperature. Adding exogenous antioxidants (D-mannitol and L-ascorbic acid) to the water surrounding the coral had no clear effect on either photoinhibition or symbiont mortality. These data show that light and temperature stress cause mortality of the dinoflagellate symbionts within the coral, and that susceptibility to light and temperature stress is strongly related to coral condition. Photoinhibitory mechanisms are clearly involved, and will increase through a positive feedback mechanism: symbiont loss promotes further symbiont loss as the light microenvironment becomes progressively harsher.