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 2.5×10⁶ to 0.8×10⁶ dinoflagellates cm² 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.     

Response of zooxanthellae in symbiosis with the Mediterranean corals Cladocora caespitosa and Oculina patagonica to elevated temperatures

Scleractinian symbiotic corals living in the Ligurian Sea (NW Mediterranean Sea) have experienced warm summers during the last decade, with temperatures rapidly increasing, within a few days, to 3–4°C above the mean value of 24°C. The effect of elevated temperatures on the photosynthetic efficiency of zooxanthellae in symbiosis with temperate corals has not been well investigated. In this study, the corals, Cladocora caespitosa and Oculina patagonica were collected in the Ligurian Sea (44°N, 9°E), maintained during 2 weeks at the mean summer temperature of 24°C and then exposed during 48 h to temperatures of 24 (control), 27, 29 and 32°C. Chlorophyll (chl) fluorescence parameters [F _v/F _m, electron transport rate (ETR), non-photochemical quenching (NPQ)] were measured using pulse amplitude modulated (PAM) fluorimetry before, during the thermal increase, and after 1 and 7 days of recovery (corals maintained at 24°C). Zooxanthellae showed a broad tolerance to temperature increase, since their density remained unchanged and there was no significant reduction in their maximum quantum yield (F _v/F _m) or ETR up to 29°C. This temperature corresponded to a 5°C increase compared to the mean summer temperature (24°C) in the Ligurian Sea. At 32°C, there was a significant decrease in chl contents for both corals. This decrease was due to a reduction in the chl/zooxanthellae content. For C. caespitosa, there was also a decrease in ETR_max, not associated with a change in F _v/F _m or in the non-photochemical quenching (NPQ); for O. patagonica, both ETR_max and F _v/F _m significantly decreased, and NPQ_max showed a significant increase. Damages to the photosystem II appeared to be reversible in both corals, since F _v/F _m values returned to normal after 1 day at 24°C. Zooxanthellae in symbiosis with the Mediterranean corals investigated can therefore be considered as resistant to short-term increases in temperature, even well above the maximum temperatures experienced by these corals in summer.     

Is acclimation beneficial to scleractinian corals,Porites spp.?

With coral reefs impacted by climate change, attention is turning to the extent to which scleractinian corals can acclimatize to new physical conditions. The implicit assumption that acclimatization is beneficial has not been tested for scleractinians, although it has been investigated in other systems and has been referred to as the beneficial acclimation hypothesis (BAH). This study tests this hypothesis for scleractinians in experiments on massive Porites spp. from Moorea, French Polynesia (17° 28.564S, 149° 49.018 W). Corals were acclimated for 15–21 days to three temperatures within the range experienced in the collection habitat and then transferred to each of the same temperatures for a treatment period of 14–15 days. The response of the holobiont was measured as growth, the response of the Symbiodinium populations as maximum photochemical efficiency of open reaction centers II (F _v/F _m). An ANOVA with polynomial contrasts was used to distinguish among the BAH, three alternative hypotheses and a null hypothesis describing the consequences of acclimation. In the first experiment (2009), massive Porites spp. were unresponsive to temperature. In the second experiment (2013), the BAH was not supported, but growth of the holobiont conformed to the “hotter is better” (HIB) hypothesis; the response of Symbiodinium populations conformed to developmental buffering. These results suggest that acclimation by massive Porites spp. to temperatures experienced routinely in the natural environment does not have clear beneficial value for growth or photochemical efficiency (i.e., BAH was not supported), but they reveal that acclimation to increased temperature can have value in responding to a variety of subsequent temperatures (i.e., support for HIB).     

Effect of acclimatization to low temperature and reduced light on the response of reef corals to elevated temperature

This study tested the effects of acclimatization on the response of corals to elevated temperature, using juvenile massive Porites spp. and branching P. irregularis from Moorea (W149°50′, S17°30′). During April and May 2006, corals were acclimatized for 15 days to cool (25.7°C) or ambient (27.7°C) temperature, under shaded (352 μmol photons m⁻² s⁻¹) or ambient (554 μmol photons m⁻² s⁻¹) natural light, and then incubated for 7 days at ambient or high temperature (31.1°C), under ambient light (659 μmol photons m⁻² s⁻¹). The response to acclimatization was assessed as biomass, maximum dark-adapted quantum yield of PSII (F _v/F _m), and growth, and the effect of the subsequent treatment was assessed as F _v/F _m and growth. Relative to the controls (i.e., ambient temperature/ambient light), massive Porites spp. responded to acclimatization through increases in biomass under ambient temperature/shade, and low temperature/ambient light, whereas P. irregularis responded through reduced growth under ambient temperature/shade, and low temperature/ambient light. Acclimatization affected the response to thermal stress for massive Porites spp. (but not P. irregularis), with an interaction between the acclimatization and subsequent treatments for growth. This interaction resulted from a lessening of the negative effects of high temperature after acclimatizing to ambient temperature/shade, but an accentuation of the effect after acclimatizing to low temperature/shade. It is possible that changes in biomass for massive Porites spp. are important in modulating the response to high temperature, with the taxonomic variation in this effect potentially resulting from differences in morphology. These results demonstrate that corals can acclimatize during short exposures to downward excursions in temperature and light, which subsequently affects their response to thermal stress. Moreover, even con-generic taxa differ in this capacity, which could affect coral community structure. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effect of sub-lethal damage to juvenile colonies of massive Porites spp. under contrasting regimes of temperature and water flow

In this study, juvenile colonies of massive Porites spp. (a combination of P. lutea and P. lobata) from the lagoon of Moorea (W 149°50′, S 17°30′) were damaged and exposed to contrasting conditions of temperature and flow to evaluate how damage and abiotic conditions interact to affect growth, physiological performance, and recovery. The experiment was conducted in April and May 2008 and consisted of two treatments in which corals were either undamaged (controls) or damaged through gouging of tissue and skeleton in a discrete spot mimicking the effects of corallivorous fishes that utilize an excavating feeding mode. The two groups of corals were incubated for 10 days in microcosms that crossed levels of temperature (26.7 and 29.6°C) and flow (6 and 21 cm s⁻¹), and the response assessed as overall colony growth (change in weight), dark-adapted quantum yield of PSII (F _v/F _m), and healing of the gouged areas. The influence of damage on growth was affected by temperature, but not by flow. When averaged across flow treatments, damage promoted growth by 25% at 26.7°C, but caused a 25% inhibition at 29.6°C. The damage also affected F _v/F _m in a pattern that differed between flow speeds, with a 10% reduction at 6 cm s⁻¹, but a 4% increase at 21 cm s⁻¹. Regardless of damage, F _v/F _m at 21 cm s⁻¹ was 11% lower at 26.7°C than at 29.6°C, but was unaffected by temperature at 6 cm s⁻¹. The lesions declined in area at similar rates (4–5% day⁻¹) under all conditions, although the tissue within them regained a normal appearance most rapidly at 26.7°C and 6 cm s⁻¹. These findings show that the response of poritid corals to sub-lethal damage is dependent partly on abiotic conditions, and they are consistent with the hypothesis that following damage, calcification and photosynthesis can compete for metabolites necessary for repair, with the outcome affected by flow-mediated mass transfer. These results may shed light upon the ways in which poritid corals respond to biting by certain corallivorous fishes.     

Ammonium enhancement of dark carbon fixation and nitrogen limitation in zooxanthellae symbiotic with the reef coralsMadracis mirabilis andMontastrea annularis

The nutrient status (limitation vs sufficiency) of dinoflagellates (zooxanthellae) symbiotic with reef corals in Bermuda was assessed in 1989 and 1990 by measuring the enhancement of dark carbon fixation with 20 M ammonium by isolated symbionts. A colony ofMadracis mirabilis was kept in the laboratory and fed daily or starved for one month. Symbionts from fed portions of the colony had ammonium-enhancement ratios (NH _4dark ⁺ ; SW_dark;SW=seawater without added ammonium) similar to those of the original field population (1.2 to 1.3). Ammonium-enhancement ratios increased with starvation of the host (x1.7) as did values forV _D:V _L [(ammonium dark rate-seawater dark rate): light rate in seawater]. Both parameters indicated decreasing nitrogen sufficiency of the algae when the host was not fed, but starvation appeared to affect these algae less than symbionts of sea anemones. Field samples of zooxanthellae fromM. mirabilis (Three Hill Shoals and Bailey's Bay Flats) yielded results similar to those for fed corals, but those taken from Bailey's Bay Flats in May 1990 yielded exceptionally high values for enhancement (>3) andV _D:V _L indicating pronounced nitrogen limitation at the time of sampling. We sampled zooxanthellae from populations ofMontastrea annularis at 8 m (Three Hill Shoals) and 24 m (Soldier's Point) depths. Enhancement andV _D:V _L values for zooxanthellae from the 8 m corals were density-dependent: symbionts from corals with normal symbiont densities displayed the most nitrogen limitation (enhancement values=1.4 to 2.0), while those from bleached corals with lower density exhibited enhancement andV _D:V _L values typical of nitrogen-sufficient algae. Symbionts isolated from the 25 m corals yielded the highest values, and appeared to exhibit the least nitrogen-sufficiency for this species.     

Effects of temperature on the mortality and growth of Hawaiian reef corals

Three common species of Hawaiian reef corals, Pocillopora damicornis (L.), Montipora verrucosa (Lamarck) and Fungia scutaria Lamarck, were grown in a temperature-regulated, continuous-flow sea water system. The skeletal growth optimum occurred near 26°C, coinciding with the natural summer ambient temperature in Hawaii, and was lowest at 21° to 22°C, representing Hawaiian winter ambient. Levels of approximately 32°C produced mortality within days. Prolonged exposure to temperatures of approximately 30°C eventually caused loss of photosynthetic pigment, increased mortality, and reduced calcification. Corals lived only 1 to 2 weeks at 18°C. The corals showed greater initial resistance at the lower lethal limit, but ultimately low temperature was more deleterious than high temperature. Results suggest that a decrease in the natural water temperature of Hawaiian reefs would be more harmful to corals than a temperature increase of the same magnitude.Contribution No. 504 of the Hawaii Institute of Marine Biology.     

Effects of ambient levels of solar ultraviolet radiation on zooxanthellae and photosynthesis of the reef coral Montipora verrucosa

Paired flat plates of the hermatypic coral Montipora verrucosa from Kaneohe Bay, Oahu, Hawaii, were acclimated to photosynthetically active radiation (PAR) only and to full sunlight (PAR+UV) for several weeks in the summer of 1990. After the acclimation period, photosynthesis, both in PAR-only and PAR+UV as well as dark respiration were measured. Levels of the UV-absorbing compounds, S320, density of zooxanthellae, and chlorophyll a concentration were determined. Corals acclimated in PAR+UV had higher levels of the UV-protective compounds and lower areal zooxanthellae densities than corals acclimated in PAR-only. Chlorophyll a per unit volume of coral host and per algal cell did not differ between corals from the two acclimation treatments. Corals acclimated to PAR+UV displayed higher photosynthesis in full sunlight than corals acclimated to PAR-only, but when photosynthesis was measured in the light regime to which the corals had been acclimated, there were no differences in photosynthesis. Dark respiration was the same for corals from the two acclimation treatments regardless of the light quality immediately preceding the dark period.Contribution No. 902 HIMB     

Monthly skeletal extension rates of the hermatypic corals<Emphasis Type="Italic"> Montastraea annularis</Emphasis> and<Emphasis Type="Italic"> Montastraea faveolata</Emphasis>: biological and environmental controls

Monthly skeletal extension rates were measured in colonies of Montastraea annularis and M. faveolata growing at Mahahual and Chinchorro Bank, in the Mexican Caribbean. Temperature, light extinction coefficient (k_d), sedimentation rate, dissolved nutrients and wave energy were used as indicators of environmental conditions for coral growth. Zooxanthella density and mitotic index, nitrogen, phosphorous and protein in coral tissue, and living tissue thickness were measured during periods of high-density-band (HDB) and low-density-band (LDB) formation. To test their value as indirect measures of competition between zooxanthellae and host, as well as coral health and performance in both species, these biological parameters were also measured, during the HDB-formation period, in corals collected at La Blanquilla. This reef is located in the Gulf of Mexico, in an area of suboptimal environmental conditions for coral growth. M. faveolata had a significantly higher skeletal extension rate than M. annularis. Corals growing in Mahahual had significantly higher skeletal extension rate than those living in Chinchorro Bank. This is consistent with inshore–offshore gradients in growth rates observed by other authors in the same and other coral species. This is probably due to less favorable environmental conditions for coral growth in near shore Mahahual, where there is high hydraulic energy and high sedimentation rate. Contrary to observations of other authors, skeletal extension rate did not differ significantly between HDB- and LDB-formation periods for both species of Montastraea. Both species produced their HDB between July and September, when the seawater temperatures are seasonally higher in the Mexican Caribbean. Tissue thickness indicated that environmental conditions are more favorable for coral health and performance during the HDB-formation period. Mitotic index data support the idea that zooxanthellae have competitive advantages for carbon over the host during the LDB-formation period. So, corals, during the LDB-formation period, with less favorable environmental conditions for coral performance and at a disadvantage for carbon with zooxanthellae, add new skeleton with little or no opportunity for thickening the existing one. This results in an equally extended skeleton with lower density, and the stretching response of skeletal growth, proposed for M. annularis growing under harsher environmental conditions, also occurs during the LDB-formation period.Communicated by P.W. Sammarco, Chauvin     

The effects of an abnormal decrease in temperature on the Eastern Pacific reef-building coral Pocillopora verrucosa

Coral bleaching events are associated with abnormal increases in temperature, such as those produced during El Niño. Recently, a breakdown in the coral–dinoflagellate (genus Symbiodinium) endosymbiosis has been documented in corals exposed to anomalously cold-water temperatures associated with La Niña events. Given the ecological significance of such events, as well as the threat of global climate change, surprisingly little is known about the physiological response of corals to cold stress. This study evaluated some physiological effects of continuous temperature decline in colonies of the eastern Pacific reef-building coral Pocillopora verrucosa. Twenty days of incubation at 18.5–19.0 °C resulted in a substantial decrease in holobiont lipid and Chla content, as well as an increase in Symbiodinium density. These observations suggest a combination of symbiont acclimation due to the temperature decline and reallocation of carbon toward algal growth as opposed to translocation to the host coral. With a decreased availability of symbiont-derived carbon, the coral likely catabolized storage lipids in order to survive the stress event. Despite this stress and some tissue necrosis, no mortality was noted and corals recovered quickly when returned to the ambient temperature. As these results are in marked contrast to similar studies investigating elevated temperature on this coral from this same location, Pocillopora in the Mexican Central Pacific may be more prone to long-term damage and mortality during periods of ocean warming as opposed to ocean cooling.     

Association of <Emphasis Type="Italic">Waminoa</Emphasis> sp. (Acoela) with corals in the Wakatobi Marine Park,South-East Sulawesi,Indonesia

This is the first quantitative study on the prevalence of epizoic Waminoa sp. acoel worms and their association with corals in the Wakatobi Marine National Park (WMNP), South-East Sulawesi, Indonesia. Three replicate transects were laid on the reef crest, flat and slope at six sites in 2006 and eight sites in 2007. Four of the sites were common in both years. In total 69 transects were surveyed in 2006, and 87 transects in 2007. A total of 4.8% of all observed hard corals were associated with acoel worms in 2006 and 2.6% of hard and soft corals in 2007. Acoels were present on 16 and 21 of the coral taxa studied in 2006 and 2007 respectively. The worms were strongly associated with the azooxanthellate coral Tubastrea spp. and were rare or absent on the most abundant coral genera Montipora and Porites. The mean number of corals having acoels was highest on reef slopes, whereas acoels were virtually absent on reef flats. Corals that had a high and a medium cover of worms were more common in 2007 than in 2006. No significant trend in the adaptation of the zooxanthellae of Waminoa sp. to different depths at different sites was revealed. The impact of the worm on the coral is unknown, but high numbers may have a shading effect and a negative impact on the coral’s photophysiology. This acoel merits more study of its life cycle, its photophysiology, and its impact on its host corals. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Improved survival under heat stress in intertidal embryos (<Emphasis Type="Italic">Fucus</Emphasis> spp.) simultaneously exposed to hypersalinity and the effect of parental thermal history

Intertidal organisms exposed to thermal stress normally experience other stresses simultaneously, but how these combined stresses modify tolerance to heat, especially for embryos, is poorly understood. Tolerance of fucoid algal embryos to heat, with and without acclimation to a sublethal temperature and with simultaneous exposure to hypersaline media, was examined. Embryos of Fucus vesiculosus L. (mid-intertidal zone) were less tolerant than embryos of Fucus spiralis L. (upper intertidal zone); without acclimation and with a growth temperature of 14°C, about half of the embryos survived 3 h exposure to 33°C in F. vesiculosus and of 35°C in F. spiralis. Conditions experienced by parental thalli (4°C versus 14°C storage) significantly affected the heat tolerance of embryos grown for 24 h post-fertilization at 14°C in F. vesiculosus, a result that is important for biologists using fucoid algae as model systems. Acclimation to a sublethal temperature (29°C) or exposure to the LT₅₀ (33°C, F. vesiculosus; 35°C, F. spiralis) in 100 psu seawater (2850 mmol kg^–1 osmolality) resulted in 30–50% higher levels of embryonic survival. Higher levels of HSP60s were found in embryos exposed to 29–33°C than to 14°C; lower levels of HSP60s were present in embryos exposed to the LT₅₀ under hypersaline conditions than in normal seawater. Contemporaneous studies in 1995–1996 of substratum temperature and desiccation levels were made at Schoodic Point, Maine (USA) underneath F. spiralis and F. vesiculosus canopies and in Semibalanus balanoides patches. This study extends the bioindicator utility of heat-shock proteins in studies of intertidal organisms and demonstrates the importance of integrated stress responses in survival of a single stress factor (e.g. temperature).Electronic Supplementary Material Supplementary material is available in the online version of this article at     

Effects of sheltering fish on growth of their host corals

Stony corals are the foundation species of tropical reefs, and their structures can harbor a diverse range of mutualist taxa that can confer important benefits, including provision of nutrients. Prominent among the associates of branching coral in the genus Pocillopora are groups of zooplanktivorous damselfishes that take refuge in the coral to avoid their predators. In field and laboratory experiments, we explored the effects of colonies of resident damselfishes on growth of their host corals. Laboratory studies revealed a positive relationship between biomass of fish and output of ammonium. In the field, levels of ammonium were significantly elevated in the water surrounding the branches of Pocillopora occupied by colonies of damselfish, particularly in time periods following active feeding by the fish. Experimental manipulation of the presence of fish on host corals during a month-long field experiment revealed that corals hosting fish grew significantly more than those that lacked fish, and coral growth was positively correlated with the biomass of resident fish. The Pocillopora colonies in the field experiment varied in the degree of openness of their branching structure, and dye studies indicated that this affected their ability to retain waterborne nutrients. Together with biomass of resident fish, colony openness explained 76% of the variation in coral growth rate during the experiment. Corals can exhibit considerable morphological variability, and mutualistic fish respond to colony architecture during habitat selection, with some species preferring more open-branched forms. This makes it likely that corals may face tradeoffs in attracting resident fish and in retaining the nutrients they provide.     

Surface-film microbial populations: diel amino acid metabolism,carbon utilization,and growth rates

Microheterotrophic dissolved free amino acid (DFAA) utilization, and microbial community and bacterial community carbon production and growth were studied using ³H-labeled organics as tracers in marine surface-film and subsurface (10 cm) waters off Baja California in November 1983. DFAA utilization was generally more rapid during the day (0.14 to 0.38 nM h^-1) than at night (0.04 to 0.14 nM h^-1) in surface-film and subsurface waters, but the percent of utilized amino acid which was respired was always greater during the night (22 to 57%) compared to the day (14 to 18%). Utilization of DFAA-carbon was estimated to range from 0.3 to 5.3 g C l^-1 d^-1 for all stations studied. In six of the 8 samples examined, the percentage of microbial carbon accounted for by the bacterial component of the population (1.4 to 5.9%) was strikingly similar to the percentage of microbial carbon production accounted for by bacterial carbon production (1.9 to 5.1%). In all of these six samples, total microbial specific-growth rates and bacterial specific-growth rates were approximately equivalent (0.9 to 2.2 d^-1 for the microbial community; 0.7 to 1.9 d^-1 for bacteria). The two exceptions were samples apparently influenced by transient flagellate populations migrating into the surface or subsurface waters at night. These observations support the conclusion that surface films contain unique and highly active microbial populations.     

Effects of particulate peat on the behavior and physiology of the Jamaican reef-building coral Montastrea annularis

Corals in an in situ respirometer exposed to suspended peat during the day greatly decreased net oxygen production, probably due to a reduction of intensity and spectral quality of light reaching the symbiotic zooxanthellae. Net production returned to pre-exposure levels after the chambers were cleared; the corals showed no behavioral effects. In contrast, after exposure during the night, corals displayed clearing behavior (such as extreme distension of the coenosarc and trapping of peat particles in thick clumps of mucus) and an increase in respiration rate comparable to the decrease in net production observed during the daytime exposure. The following morning, net production values were significantly lower than pre-exposure production values although ambient light intensity was slightly higher. This decrease in production as well as a 22% reduction of chlorophyll content in the coral tissue indicated loss of zooxanthellae from the stressed corals. Long-term exposure to such a stress could reduce coral growth rates and substantially alter coral reef communities.     

Effect of hyposalinity on the photophysiology of <Emphasis Type="Italic">Siderastrea radians</Emphasis>

Tolerance to hyposalinity of the scleractinian coral S. radians was examined in a mesocosm study. Colonies of S. radians were collected from five basins in Florida Bay, USA, which occur along a northeast-to-southwest salinity gradient. Salinity treatments were based on historical salinity records for these basins. Photophysiology of the endosymbiont Symbiodinium spp. (maximum quantum yield; F _v/F _m) was measured as an indicator of holobiont stress to hyposalinity. Colonies from each basin were assigned four salinity treatments [The Practical Salinity Scale (PSS) was used to determine salinity. Units are not assigned to salinity values because it is a ratio and has no unit as defined by UNESCO (UNESCO Technical papers no. 45, IAPSO Pub. Sci. No. 32, Paris, France, 1985)] (30, 20, 15, and 10) and salinities were reduced 2 per day from ambient (30) to simulate a natural salinity decrease. Colonies treated with salinities of 20 and 15 showed no decrease in F _v/F _m versus controls (i.e. 30), up to 5 days after reaching their target salinity. This indicates a greater ability to withstand reduced salinity for relatively extended periods of time in S. radians compared to other reef species. Within 1 day after salinity of 10 was reached, there was a significant reduction in F _v/F _m, indicating a critical threshold for hyposaline tolerance. At the lowest treatment salinity (10), F _v/F _m for the more estuarine, northeast-basin colonies were significantly higher than the most marine southwest-basin colonies (Twin Key Basin). Our results suggest that historical salinity ranges within basins determine coral population salinity tolerances.     

Influence of light on algal symbionts of the deep water coral Leptoseris fragilis

Photoadaptations of zooxanthellae living within the deep water coral Leptoseris fragilis taken from the Gulf of Aqaba (Red Sea) were studied. Specimens-collected in summer 1988 between 110 and 120 m depth —were transplanted to 70 and 160 m. At each depth individuals were exposed in their natural growth position (oral side facing the surface) or in a reverse growth position (oral side facing the bottom). After 1 yr of exposure the corals were collected and the zooxanthellae were isolated. As a function of the availability of light with depth and growth position several algal parameters showed changes which are related to photoadaptations. The relatively low density of zooxanthellae of 0.15x10⁶ cellsxcm^-2 at a natural growth depth of 116 m decreased to 0.0034x10⁶ cellsxcm^-2 (2%) at 160 m in specimens growing with a natural orientation. In corals with a downward-facing oral surface at the same depth (160 m) only degenerated algae could be observed. With respect to depth dependence the volume of the algae decreased from 728 m³ at 116 m to 406 m³ at a depth of 160 m and the content of pigments increased. The augmentation of peridinin per cell was low (two times at 160 m compared to 116 m). Chlorophyll a and in particular chlorophyll c ₂ concentrations per cell were enhanced. Compared to natural amounts at 116 m, chl a was five times and chl c ₂ eight times higher at 160 m. At all depths the chl c ₂ content per cell was higher than for chl a. The formation of chl a/chl c ₂ complexes as light harvestor is discussed. Light harvesting, with chl c ₂ prevailing may be explained as a special type of chromatic adaptation of L. fragilis in a double sense: (1) in the habitat light short wavelengths predominate. This light can be directly absorbed with pigments such as chl a and chl c ₂. (2) Host pigments absorb visible violet light and transform these wavelengths, less suitable for photosynthesis, into longer ones by means of autofluorescence. The emitted longer wavelengths fit the absorption maxima of the algal pigments. Thus the host supports photosynthesis of his symbionts. Corals exposed at 160 m depth with a downward facing oral surface were alive after 1 yr and the host wavelength transforming pigment system was still present, but zooxanthellae were absent or degenerated. The light field at 160 m seems therefore to be critical: the combined photoadaptations of host and symbionts, allowing photosynthesis under barren light conditions, seem to be exhausted. In L. fragilis the photoadaptive strategies of host and symbionts cooperate harmoniously. In addition, the adaptations are interlocked with the particular light situation of the habitat with respect to light quantity and quality. The cooperation of physical and organismic parameters examplifies how evolution and, in particular, coevolution has led to optimal fitness.     

Stability and Mixing of a Vertical Axisymmetric Buoyant Jet in Shallow Water

The stability, mixing and effect of downstream control on axisymmetric turbulent buoyant jets discharging vertically into shallow stagnant water is studied using 3D Reynolds-averaged Navier–Stokes equations (RANS) combined with a buoyancy-extended k –ε model. The steady axisymmetric turbulent flow, temperature (or tracer concentration) and turbulence fields are computed using the finite volume method on a high resolution grid. The numerical predictions demonstrate two generic flow patterns for different turbulent heated jet discharges and environmental parameters (i) a stable buoyant discharge with the mixed fluid leaving the vertical jet region in a surface warm water layer; and (ii) an unstable buoyant discharge with flow recirculation and re-entrainment of heated water. A stratified counterflow region always appears in the far-field for both stable and unstable buoyant discharges. Provided that the domain radius L exceeds about 6H, the near field interaction and hence discharge stability is governed chiefly by the jet momentum length scale to depth ratio l_M/H, regardless of downstream control. The near field jet stability criterion is determined to be l_M/H = 3.5. A radial internal hydraulic jump always exists beyond the surface impingement region, with a 3- to 6-fold increase in dilution across the jump compared with vertical buoyant jet mixing. The predicted stability category, velocity and temperature/concentration fields are well-supported by experiments of all previous investigators.     

Laboratory observations on spawning,fecundity and early development of a mesopelagic ostracod,Conchoecia pseudodiscophora,from the Japan Sea

The fate of the polycyclic aromatic hydrocarbon [³H]benzo[a]pyrene (BaP) was examined in two species of scleractinian corals, Favia fragum (Esper) and Montastrea annularis (Ellis and Solander), which were collected in the patch reefs surrounding Alina's Reef (25°23.25N; 80°09.8W) in Biscayne National Park, Florida, USA, in July, 1990. Corals were exposed to initial concentrations of 5 g/l in a simple static system for 25 h. BaP uptake was estimated from the disappearance of BaP from the water. Uptake rates were 6.5±0.7 and 10.8±0.2 g BaP cm^-2h^-1 for F. fragum and M. annularis, respectively, at initial BaP concentrations and were directly proportional to the concentration of BaP in the water. The separation of zooxanthellae from coral tissue revealed that zooxanthellae can accumulate up to 53 and 64% of the total BaP-derived radioactivity present in F. fragum and M. annularis, respectively. Both corals metabolized BaP slowly, as most of the accumulated radioactivity was present as the unmetabolized chemical. However, aqueous and organic-soluble metabolites were found in both the animal and zooxanthellae fractions. Analysis by high-performance liquid chromatography (HPLC) revealed that both species of corals metabolized BaP to various tetrols, triols, dihydrodiols, quinones and phenols, although the pattern of metabolites differed between species. Zooxanthellae contained some of the same Phase I metabolites found in the animal tissue; however, tetrols and triols were absent in extracts from the zooxanthellae. The elimination of BAP from corals was also slow; approximately 38 and 65% of the accumulated radioactivity was still present in F. fragum and M. annularis, respectively, 144 h following the transfer of exposed corals to an uncontaminated flow-through seawater system.     

Phototaxis in Fungiidae corals (Scleractinia)

Corals within the scleractinian family Fungiidae were observed to move toward light (positive phototaxis). Negative phototactic movement was not observed in any of the specimens tested. On sandy substratum, Diaseris distorta moved faster (max. speed of 3 cm h^-1) than other species. Although D. distorta has symbiotic algae, phototactic movement also was observed both in bleached corals and in those treated with a specific inhibitor (dichlorophenyl dimethyl urea) of photosynthesis. D. distorta was phototactic even on a glass plate, and climbed up a steep slope (up to 30°). Based on experiments with Fungia fungites and D. distorta, soft tissues at the peripheral region of the dise seem to be responsible for movement via peristalsis. It is suggested that positive phototaxis in symbiontbearing fungiid corals is an important trait for selection of favorable habitats.