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

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

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

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

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

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

Spring “bleaching” among <Emphasis Type="Italic">Pocillopora</Emphasis> in the Sea of Cortez,Eastern Pacific

A mild bleaching event was observed among Pocillopora spp. in the southern Gulf of California in the spring of 2006. Uniform bleaching occurred in numerous colonies on the upper portions of their branches. Most (∼90%) colonies that exhibited bleaching contained a species of endosymbiotic dinoflagellate, Symbiodinium C1b-c, which differed from the Symbiodinium D1 found inhabiting most unbleached colonies. Analysis of chlorophyll fluorescence, indicated a decline in photosystem II photochemical activity, especially among colonies populated with C1b-c. By early August, most affected colonies had recovered their normal pigmentation and fluorescence values were once again high for all colonies. No mortality was observed among tagged bleached colonies nor did symbiont species composition change during recovery. This unusual episode of bleaching did not appear to be a response to thermal stress, but may have been triggered by high levels of solar radiation during a period of unseasonally high water clarity in the early spring.     

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

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

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

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

Stress response of two coral species in the Kavaratti atoll of the Lakshadweep Archipelago, India

Frequent occurrences of coral bleaching and the ensuing damage to coral reefs have generated interest in documenting stress responses that precede bleaching. The objective of this study was to assess and compare physiological changes in healthy, semi-bleached and totally bleached colonies of two coral species, Porites lutea and Acropora formosa, during a natural bleaching event in the Lakshadweep Archipelago in the Arabian Sea to determine the traits that will be useful in the diagnosis of coral health. In April 2002, three “health conditions” were observed as “appearing healthy,” “semi-bleached” and “bleached” specimens for two dominant and co-occurring coral species in these islands. Changes in the pigment composition, zooxanthellae density (ZD), mitotic index (MI) of zooxanthellae, RNA/DNA ratios and protein profile in the two coral species showing different levels of bleaching in the field were compared to address the hypothesis of no difference in health condition between species and bleaching status. The loss in chlorophyll (chl) a, chl c and ZD in the transitional stage of semi-bleaching in the branched coral A. formosa was 80, 75 and 80%, respectively. The losses were much less in the massive coral P. lutea, being 20, 50 and 25%, respectively. The decrease in zooxanthellar density and chl a was accompanied by an increased MI of zooxanthellae and RNA/DNA ratios in both the species. There was an increase in accumulation of lipofuscin granules in partially bleached P. lutea tissue, which is an indication of cellular senescence. Multivariate statistical analyses showed that colonies of P. lutea ranked in different health conditions differed significantly in chl a, chl c, ZD, RNA/DNA ratios, and protein concentrations, whereas in A. formosa chl a, chl c, chl a/c, phaeopigments and MI contributed to the variance between health conditions.     

Detrimental effects of host anemone bleaching on anemonefish populations

Coral bleaching and related reef degradation have caused significant declines in the abundance of reef-associated fishes. Most attention on the effects of bleaching has focused on corals, but bleaching is also prevalent in other cnidarians, including sea anemones. The consequences of anemone bleaching are unknown, and the demographic effects of bleaching on associated fish recruitment, survival, and reproduction are poorly understood. We examined the effect of habitat degradation including host anemone bleaching on fish abundance, egg production, and recruitment of the panda anemonefish (Amphiprion polymnus) near Port Moresby, Papua New Guinea. Following a high-temperature anomaly in shallow waters of the region, most shallow anemones to a depth of 6 m (approximately 35% of all the anemones in this area) were severely bleached. Anemone mortality was low but bleached anemones underwent a ~34% reduction in body size. Total numbers of A. polymnus were not affected by bleaching and reduction in shelter area. While egg production of females living in bleached anemones was reduced by ~38% in 2009 compared to 2008, egg production of females on unbleached anemones did not differ significantly between years. Total recruitment in 2009 was much lower than in 2008. However, we found no evidence of recruiting larvae avoiding bleached anemones at settlement suggesting that other factors or different chemical cues were more important in determining recruitment than habitat quality. These results provide the first field evidence of detrimental effects of climate-induced bleaching and habitat degradation on reproduction and recruitment of anemonefish.     

Cellular processes of bleaching in the Mediterranean coral Oculina patagonica

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

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

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

Microscopic observations of recovery in the reef-building scleractinian coral,Montastrea annularis,after bleaching on a Cayman reef

Living tissues of a single massive colony of the reef-building scleractinian coral, M. annularis, were sampled at one month intervals following the observation that areas of marked discoloration (bleaching) had developed by the beginning of the summer of 1988. Histological analysis of replicate plugs of tissue from bleached and unbleached regions of that colony allowed comparison of time-matched samples as well as zone-matched samples over a period of six months. During that interval, the entire colony was restored to a uniform and normal degree of pigmentation. Histology of the bleached zones documents the gradual accumulation of zooxanthellae and the thickening of the gastrodermal epithelium of the coral. Once reconstituted, the previously bleached gastrodermis, repopulated with algal symbionts, is indistinguishable from unbleached tissue in the coral colony.     

Reduced growth rate of Montastrea annularis following the 1987–1988 coral-bleaching event

Montastrea annularis, the major Caribbean reef building coral, was severely affected by the unprecedented 1987–1988 bleaching event. Most colonies on the fore reef were affected but few were bleached in the back reef. Skeletal growth rates of M. annularis populations were measured non-destructively in the field at Discovery Bay, Jamaica, from the peak of bleaching in Nov. 1987 until recovery was almost complete, in May 1988. Unbleached corals grew at normal rates. Partially bleached corals survived but skeletal growth ceased through this period.     

Specific bleaching of phycobiliproteins from cyanobacteria and red algae at high temperature in vivo

Exposure of blue-green or red algal cells to temperatures exceeding 60–65°C for several minutes resulted in bleaching of all phycobilin absorption in the visible range, with virtually no alteration in chlorophyll or carotenoid absorption. Difference spectra of non-bleached vs bleached cells appeared identical to absorption spectra of purified phycobilisomes isolated from the same cell culture in high phosphate medium. All phycobilin chromophores were bleached at approximately the same rate during heating. There were no changes in apparent molecular weights or relative amounts of the phycobilisome apoproteins during chromophore bleaching. Phycobilisomes in cell extracts from Anacystis nidulans resisted bleaching when suspended in medium of high phosphate concentration, but were bleached at 60–65°C within a few minutes when placed in diluted medium. The results indicate that phycobilisomes in vivo are stabilized by a mechanism other than high osmotic and ionic strength. This represents a rapid and quantitative method to characterize the phycobiliprotein content of cyanobacteria and red algae in vivo.Abbreviations Chl chlorophyll - APC allophycocyanin - PC phycocyanin - PE phycoerythrin - SPM medium, 0.2 M sucrose, 15 mM MgCl₂, 0.75 M Na/KPO₄, pH 7.8     

Estimating the Size of HIV Key Affected Populations in Chongqing,China, Using the Network Scale-Up Method

Objectives

To estimate the average social network size in the general population and the size of HIV key affected populations (KAPs) in Chongqing municipality using the network scale-up method (NSUM).

Methods

A general population survey was conducted in 2011 through a multistage random sampling method. Participants aged between 18 and 60 years were recruited. The average social network size (c) was estimated and adjusted by known population method. The size of HIV KAP in Chongqing municipality was estimated using the adjusted c value with adjustment for the transmission effect using the scaled respect factor.

Results

3,026 inhabitants of Chongqing agreed to the survey, and 2,957 (97.7%) completed the questionnaire. The adjusted c value was 310. The estimated size of KAP was 28,418(95% Confidence Interval (CI):26,636∼30,201) for female sex workers (FSW), 163,199(95%CI:156,490∼169,908) for clients of FSW, 37,959(95%CI: 34,888∼41,030) for drug users (DU), 14,975(95%CI:13,047∼16,904) for injecting drug users (IDU) and 16,767(95%CI:14,602∼18,932) for men who have sex with men (MSM). The ratio of clients to FSW was 5.74∶1, and IDU accounted for 39.5% of the DU population. The estimates suggest that FSW account for 0.37% of the female population aged 15–49 years in Chongqing, and clients of FSW and MSM represent 2.09% and 0.21% of the male population aged 15–49 years in the city, respectively.

Conclusion

NSUM provides reasonable population size estimates for FSW, their clients, DU and IDU in Chongqing. However, it is likely to underestimate the population size of MSM even after adjusting for the transmission effect.     

A comparison of the 1998 and 2002 coral bleaching events on the Great Barrier Reef: spatial correlation,patterns, and predictions

Detailed mapping of coral bleaching events provides an opportunity to examine spatial patterns in bleaching over scales of 10 s to 1,000 s of km and the spatial correlation between sea surface temperature (SST) and bleaching. We present data for two large-scale (2,000 km) bleaching events on the Great Barrier Reef (GBR): one from 1998 and another from 2002, both mapped by aerial survey methods. We examined a wide range of satellite-derived SST variables to determine which one best correlated with the observed bleaching patterns. We found that the maximum SST occurring over any 3-day period (max3d) during the bleaching season predicted bleaching better than anomaly-based SST variables and that short averaging periods (3–6 days) predicted bleaching better than longer averaging periods. Short periods of high temperature are therefore highly stressful to corals and result in highly predictable bleaching patterns. Max3d SST predicted the presence/absence of bleaching with an accuracy of 73.2%. Large-scale (GBR-wide) spatial patterns of bleaching were similar between 1998 and 2002 with more inshore reefs bleached compared to offshore reefs. Spatial change in patterns of bleaching occurred at scales of ~10 s km, indicating that reefs bleach (or not) in spatial clusters, possibly due to local weather patterns, oceanographic conditions, or both. Approximately 42% of reefs bleached to some extent in 1998 with ~18% strongly bleached, while in 2002, ~54% of reefs bleached to some extent with ~18% strongly bleached. These statistics and the fact that nearly twice as many offshore reefs bleached in 2002 compared to 1998 (41 vs. 21%, respectively) makes the 2002 event the worst bleaching event on record for the GBR. Modeling of the relationship between bleaching and max3d SST indicates that a 1 °C increase would increase the bleaching occurrence of reefs from 50% (approximate occurrence in 1998 and 2002) to 82%, while a 2 °C increase would increase the occurrence to 97% and a 3 °C increase to 100%. These results suggest that coral reefs are profoundly sensitive to even modest increases in temperature and, in the absence of acclimatization/adaptation, are likely to suffer large declines under mid-range International Panel for Climate Change predictions by 2050.

Assessing coral bleaching and recovery with a colour reference card in Watamu Marine Park,Kenya

With this study we estimated the changes in colour, bleaching and mortality of coral colonies from February to December 2007, using the colour reference card method. The study was developed in the Watamu Marine Park lagoon (Kenya), bridging the local summer when seawater temperatures were highest and coral bleaching risk was at its maximum. Seven coral genera were selected, and their colour recorded using a colour reference card (Coral Watch card). Seven different scenarios of bleaching and mortality were observed, varying among the coral genera and between two species in the genus Pocillopora. Twenty percent of the colonies bleached, of which 50% died. Only 15% of the coral that did not bleach died. Branching genera had a higher bleaching incidence than massive and sub-massive genera. Pocillopora showed the highest bleaching susceptibility, followed by Acropora, and the highest level of mortality. Of the two species of Pocillopora considered in this study, P. eydouxi showed higher bleaching and mortality levels, while P. verrucosa bleached less and experienced only partial mortality. Our results evidenced different patterns of coral bleaching and mortality which were easily and clearly detected with the colour card method during both bleaching and a post-bleaching events.     

Study of water permeability through phospholipid vesicle membranes by O NMR

Vesicle suspensions of up to 5 % egg lecithin and 2.5 % cholesterol have been found to have no effect on the NMR relaxation times of ¹⁷O from water. Addition of 1–5 mM Mn²⁺ to an equimolar vesicle suspension of egg lecithin and cholesterol permitted resolution of the free induction decay into two exponential components, a fast one arising from the external water and a slow one arising from the intravesicular fluid. From the rates of relaxation the mean life time of the water molecules within the vesicles was calculated to be 1±0.1 ms at 22°C. The size of the vesicle was estimated from electron micrographs to be about 500 Å in diameter. These data yield an equilibrium water permeability, P_w, of about 8 μs⁻¹ for the vesicle membranes. From the temperature dependence of P_w an activation energy of 12±2 kcal/mol was obtained. The longitudinal relaxation time (T₁) of water within vesicles remained the same as in pure water.     

Bleaching of reef coelenterates in the San Blas Islands,Panama

Starting in June 1983, 25 species of hermatypic corals, gorgonians, hydrocorals, anemones and zoanthids in the San Blas Islands, Panama, began showing signs of a loss of colour leading in some cases to a white bleached appearance. Histologic examination of six coral species indicated that bleaching was associated with drastic reductions in the density of zooxanthellae and with the atrophy and necrosis of the animal tissue. The severity of the bleaching varied among species and many species were unaffected. The species most extensively affected were: Agaricia spp., which became completely bleached and frequently died; Montastraea annularis which bleached and continued to survive; and Millepora spp. which bleached white but quickly regained their colouration. Shallow reefs dominated by Agaricia spp. suffered the most extensive bleaching. At one site, Pico Feo, 99% of the Agaricia (32% of the living cover) was bleached. On fore reers, which were dominated by Agaricia spp. and M. annularis, the proportion of M. annularis bleached ranged from 18 to 100% and that of Agaricia spp. from 30 to 53%. Transects at Sail Rock and House Reef were surveyed in August 1983 and January 1984. At those sites, 53% of the Agaricia cover died between August and January. The remaining living cover of Agaricia and of all other species exhibited normal colouration in January. Salinity and temperature were monitored every second day at 4 m depth between May 10 and August 28, 1983 at one of the localities. Bleaching was first observed within two weeks of a 2 °C rise in temperature which occurred in late May 1983. Temperatures remained at or above 31.5 °C for the following 3 weeks and were at or above 30 °C for an additional 4 weeks. The bleaching of corals in the San Blas was most likely due to those elevanted temperatures.     

The structure of Escherichia coli membranes studied by fluorescence measurements of lipid phase transitions

Lipid phase transitions in Escherichia coli membranes and in dispersions of the extracted lipids were studied using the negatively charged fluorescence probe 1-anilinonaphthalene-8-sulfonate (ANS⁻) and the hydrophobic fluorescence probe N-phenyl-1-naphthylamine (NPN). The fluorescence change, ΔI, at the phase transition approaches a limiting value (ΔI)_lim with increasing dye concentration. A comparison of the limiting values (Δ)_lim^NPN obtained for membranes and the lipid standard allows us to estimate the lipid fraction, ρ, in the membrane that takes part in the phase transition (ρ = 80%). The same procedure carried out with ANS⁻ yields a value of 42.5% for the lipid fraction that is accessible from the aqueous phase. These values, combined with published freeze-etching data for the particle density within the fracture plane of membranes are used to quantify the Davson-Danielli-Robertson-Benson-Singer membrane model which assumes a fluid lipid bilayer with “integral” proteins embedded in the lipid matrix and surface proteins attached to the lipid head groups. It appears that on the average one “integral” membrane protein is surrounded by about 600 lipid molecules and that about 130 of these molecules are closely coupled to the protein molecule, forming an halo in which the chain-chain interaction between the lipids is disturbed. About half of the bilayer surface is covered with proteins; part of these seem to be stacked.     

Characterization of fatty acid composition in healthy and bleached corals from Okinawa, Japan

Under bleaching conditions, corals lose their symbiotic zooxanthellae, and thus, the ability to synthesize fatty acids (FAs) from photosynthetically derived carbon. This study investigated the lipid content and FA composition in healthy and bleached corals from the Odo reef flat in Okinawa, southern Japan, following a bleaching event. It was hypothesized that the FA composition and abundance would change as algae are lost or die, and possibly microbial abundance would increase in corals as a consequence of bleaching. The lipid content and FA composition of three healthy coral species (Pavona frondifera, Acropora pulchra, and Goniastrea aspera) and of partially bleached and completely bleached colonies of P. frondifera were examined. The FA composition did not differ among healthy corals, but differed significantly among healthy, partially bleached, and completely bleached specimens of P. frondifera. Completely bleached corals contained significantly lower lipid and total FA content, as well as lower relative amounts of polyunsaturated FAs and higher relative amounts of saturated FAs, than healthy and partially bleached corals. Furthermore, there was a significantly higher relative concentration of monounsaturated FAs and odd-numbered branched FAs in completely bleached corals, indicating an increase in bacterial colonization in the bleached corals.