Certain stony corals can alternate between a calcifying colonial form and noncalcifying solitary polyps, supporting the hypothesis that corals have survived through geologic timescale periods of unfavorable calcification conditions. However, the mechanisms enabling this biological plasticity are yet to be identified. Here we show that incubation of two coral species (Pocillopora damicornis and Oculina patagonica) under reduced pH conditions (pH 7.2) simulating past ocean acidification induce tissue-specific apoptosis that leads to the dissociation of polyps from coenosarcs. This in turn leads to the breakdown of the coenosarc and, as a consequence, to loss of coloniality. Our data show that apoptosis is initiated in the polyps and that once dissociation between polyp and coenosarc terminates, apoptosis subsides. After reexposure of the resulting solitary polyps to normal pH (pH 8.2), both coral species regenerated coenosarc tissues and resumed calcification. These results indicate that regulation of coloniality is under the control of the polyp, the basic modular unit of the colony. A mechanistic explanation for several key evolutionarily important phenomena that occurred throughout coral evolution is proposed, including mechanisms that permitted species to survive the third tier of mass extinctions.apoptosis | ocean acidification | corals

Spatial patterns of marine Synechococcus diversity across ocean domains have been reported on extensively. However, much less is known of seasonal and multiannual patterns of change in Synechococcus community composition. Here we report on the genotypic diversity of Synechococcus populations in the Gulf of Aqaba, Northern Red Sea, over seven annual cycles of deep mixing and stabile stratification, using ntcA as a phylogenetic marker. Synechococcus clone libraries were dominated by clade II and XII genotypes and a total of eight different clades were identified. Inclusion of ntcA sequences from the Global Ocean Sampling database in our analyses identified members of clade XII from beyond the Gulf of Aqaba, extending its known distribution. Most of the Synechococcus diversity was attributed to members of clade II during the spring bloom, while clade III contributed significantly to diversity during summer stratification. Clade XII diversity was most prevalent in fall and winter. Clade abundances were estimated from pyrosequencing of the V6 hypervariable region of 16S rRNA. Members of clade II dominated Synechococcus communities throughout the year, whereas the less frequent genotypes showed a pattern of seasonal succession. Based on the prevailing nutritional conditions we observed that clade I members thrive at higher nutrient concentrations during winter mixing. Clades V, VI and X became apparent during the transition periods between mixing and stratification. Clade III became prominent during sumeer stratification. We propose that members of clades V, VI, and X, and clade III are Synechococcus ecotypes that are adapted to intermediate and low nutrient levels respectively. This is the first time that molecular analyses have correlated population dynamics of Synechococcus genotypes with temporal fluctuations in nutrient regimes. Since these Synechococcus genotypes are routinely observed in the Gulf of Aqaba we suggest that seasonal fluctuations in nutrient levels create temporal niches that sustain their coexistence.

Elevated seawater temperatures are associated with coral bleaching events and related mortality. Nevertheless, some coral species are able to survive bleaching and recover. The apoptotic responses associated to this ability were studied over 3 years in the coral Stylophora pistillata from the Gulf of Eilat subjected to long term thermal stress. These include caspase activity and the expression profiles of the S. pistillata caspase and Bcl-2 genes (StyCasp and StyBcl-2-like) cloned in this study. In corals exposed to thermal stress (32 or 34°C), caspase activity and the expression levels of the StyBcl-2-like gene increased over time (6–48 h) and declined to basal levels within 72 h of thermal stress. Distinct transcript levels were obtained for the StyCasp gene, with stimulated expression from 6 to 48 h of 34°C thermal stress, coinciding with the onset of bleaching. Increased cell death was detected in situ only between 6 to 48 h of stress and was limited to the gastroderm. The bleached corals survived up to one month at 32°C, and recovered back symbionts when placed at 24°C. These results point to a two-stage response in corals that withstand thermal stress: (i) the onset of apoptosis, accompanied by rapid activation of anti-oxidant/anti-apoptotic mediators that block the progression of apoptosis to other cells and (ii) acclimatization of the coral to the chronic thermal stress alongside the completion of symbiosis breakdown. Accordingly, the coral's ability to rapidly curb apoptosis appears to be the most important trait affecting the coral's thermotolerance and survival.

Corallimorpharians are evolutionarily important relatives to reef-building corals, yet little is known about their ecophysiology. We demonstrate that physiological mechanisms determine, in part, the light-dependent distributional patterns of 2 corallimorpharians, Rhodactis rhodostoma and Discosoma unguja, on coral reefs in the northern Red Sea. Field measurements of the physiological parameters related to photosynthetic activity revealed that zooxanthellae abundance, chlorophyll a concentration, maximum quantum yield (F v /F m ), and excitation pressure of Photosytem II (Q m ) varied equally for both species among 3 depths. In contrast, laboratory measurements in 3 experimental light treatments -low light (LL), medium light (ML) and high light (HL) -revealed that D. unguja was more sensitive to HL than R. rhodostoma. Genetic characterization of their endosymbiotic algae using a PCR for the internal transcribed spacer region (ITS2) of nuclear ribosomal DNA showed that both R. rhodostoma and D. unguja contain Symbiodinium ITS2-type C1 symbionts in shallow areas, but that the algal assemblage were either Type C1 or Type D1a in both species in deeper areas. In reciprocal depth, light, and temperature manipulation experiments, bleached R. rhodostoma never shuffled Symbiodinium ITS2 types, however, some R. rhodostoma in the control treatment did. In contrast, some individuals of D. unguja shuffled Symbiodinium ITS2 types both in field-reciprocal experiments and laboratory-control treatments. Both species contained UV radiation absorbing compounds over a wide-depth range. We concluded that the photoacclimation mechanisms of R. rhodostoma and D. unguja were influenced by both host-and symbiont-mediated factors. We hypothesize that a low abundance of D. unguja in shallow water was due to decreased tolerance to the high irradiance found in host factors. KEY WORDS: Photoacclimation mechanisms · Corallimorpharians · Photosynthetic parameters · ZooxanthellaeResale or republication not permitted without written consent of the publisher Mar Ecol Prog Ser 369: 115-129, 2008 combination of high atmospheric CO 2 concentrations and elevated temperatures (Berner 1992). During the early Cretaceous, multiple calcifying scleractinians became extinct (Buddemeier 1996), but many survived and radiated into considerable diversity (ChadwickFurman 1996, Stanley 2003. The surviving corallimorpharians may have lost their ability to calcify, but they maintained primary life functions, perhaps creating a means to increase fitness in a relatively acidic sea with low Mg/Ca ratios (Stanley 2003, Medina et al. 2006. Loss of skeleton in corallimorpharians may have been compensated for by enhanced physiological capabilities as a means of coping with environmental changes. Understanding more about the physiological properties of corallimorpharians offers an opportunity to uncover a latent chapter in the evolution of corals and their ability to survive immense climatic changes. While the effects of light and temperature stress on reef-buil...