Simulating light-saturation curves for photosynthesis and calcification by reef-building corals

Light-saturation curves for photosynthesis by reef-building corals have previously been simulated by three functions: the right rectangular hyperbola, a simple exponential function, and the hyperbolic tangent function. Studies of photosynthesis by other organisms have also frequently considered the application of a rectilinear function. This communication analyzes lightsaturation curves for photosynthesis by the Atlantic rose coral Manicina aerolata, the Atlantic staghorn coral Acropora cervicornis, and the Pacific staghorn coral A. formosa. It also analyzes light-saturation curves for calcification by A. cervicornis and A. formosa. This communication demonstrates that the two most accurate functions (as measured by coefficients of determination) are the simple exponential function and the hyperbolic tangent function. The hyperbolic tangent function is preferred because parameter estimates obtained with this function have narrower confidence intervals than those obtained through the application of the simple exponential function. The hyperbolic tangent function can also be used successfully to simulate light-saturation curves for light-enhanced calcification.     

Effects of temperature on photosynthesis and respiration in hermatypic corals

Photosynthesis and respiration rates of the reef corals Pocillopora damicornis (Linn.), Montipora verrucosa (Lamarck), Porites compressa Dana and Fungia scutaria Lamarck were measured under controlled temperatures. Results indicate that coral metabolism is closely adapted to ambient temperature conditions. Tropical corals measured at Enewetak, Marshall Islands, showed greater primary production compared to maintenance requirements at elevated temperatures than did subtropical varieties of the same species in Hawaii. Photosynthesis: respiration (P:R) ratios were significantly and negatively related with temperature between 18° and 31°C for all Hawaiian corals, whereas at Enewetak this ratio generally showed a curvilinear relationship for this temperature range. Extrapolations of P:R regressions on temperatures to a value of 2.0 (estimated as a minimum required for long-term functional autotrophy) coincide for Hawaiian specimens with published upper lethal temperatures. Extrapolation of P:R regressions for Enewetak specimens at temperatures above 25°C suggests lethal temperatures for these corals to be 2 to 5 C° higher than for Hawaiian corals, in good agreement with recent experimental findings. Interspecific differences in P:R temperature regressions for Hawaiian corals correlating with upper lethal temperature tolerances are described.Contribution No. 505 of the Hawaii Institute of Marine Biology.     

Interactions between morphological and physiological plasticity optimize energy acquisition in corals

Morphological plasticity in response to environmental heterogeneity may be performance enhancing or may simply result from an intrinsic instability in morphology during development. Although patterns of morphological change are well documented for numerous taxa, it is often unclear whether this plasticity enhances the performance of organisms in the habitat to which they have acclimatized. Reef-building corals are an ideal model system in which to investigate this question. We here develop a three-dimensional geometric model and present a comprehensive photosynthesis data set with experimentally calibrated photosynthesis models that predicts energy acquisition by foliose corals as a function of colony shape. This allows us to assess the extent to which changes in colony morphology along an environmental gradient track the predicted optimal colony morphologies. Our results provide strong evidence that phenotypic plasticity in foliose corals optimizes photosynthetic energy acquisition and is not simply a mechanism to increase light capture. We show that the optimal morphology is constrained at the boundaries of the environmental gradient, with non-optimal morphologies in these habitats having greatly reduced energy acquisition. However, at the center of the environmental gradient, flexibility in photophysiology allows energy acquisition to be very similar for multiple morphologies. Our results highlight the importance of phenotypic plasticity at multiple scales. Variation in overall morphology is important at niche boundaries at which conditions are consistently more stressful, whereas physiological flexibility is important in intermediate and less predictable habitats in which a rapid and reversible response to environmental fluctuations is required.     

The effect of inhibitors of photosynthesis on zooxanthellae in corals and other marine invertebrates

The effect of the selective photosynthesis inhibitors Monuron (CMU), Diuron (DCMU) and methyl viologen on intact algal-marine invertebrate symbiotic associations was studied. CMU or DCMU (5x10^-4M) completely inhibited photosynthesis, both in intact branches, and in suspensions of isolated zooxanthellae from the reef-building coral Pocillopora damicornis. The inhibitory effect was totally reversible in 1 to 3 h after removal of the inhibitor. Similar inhibition of photosynthesis occurred in 8 other marine coelenterates symbiotic with zooxanthellae, and in 1 marine gastropod symbiotic with functional chloroplasts. Neither CMU nor DCMU appeared to affect behavior of the various hosts, such as swimming, phototaxis, phototropism, photoreception, tentacle contraction, ciliary beating and locomotion. Methyl viologen, however, was ineffective in inhibiting photosynthesis in intact P. damicornis at low concentrations, and lethal to the tissues at high concentrations. These observations indicate that CMU and DCMU are potential useful tools for investigation of symbiotic associations. DCMU (5x10^-4M) also reversibly inhibited light-enhanced calcification in P. damicornis. This strongly suggests that light-enhanced calcification is largely photosynthesis dependent, and probably not dependent on some other photobiological effect.Contribution No. 385, Hawaii Institute of Marine Biology, University of Hawaii     

Spatial heterogeneity of photosynthesis and the effect of temperature-induced bleaching conditions in three species of corals

Heterogeneity in photosynthetic performance between polyp and coenosarc tissue in corals was shown using a new variable fluorescence imaging system (Imaging-PAM) with three species of coral, Acropora nobilis, Cyphastrea serailia and Pocillopora damicornis. In comparison to earlier studies with fibre-optic microprobes for fluorescence analysis, the Imaging-PAM enables greater accuracy by allowing different tissues to be better defined and by providing many more data points within a given time. Spatial variability of photosynthetic performance from the tip to the distal parts was revealed in one species of branching coral, A. nobilis. The effect of bleaching conditions (33°C vs. 27°C) was studied over a period of 8 h. Marked changes in fluorescence parameters were observed for all three species. Although a decline in _PSII (effective quantum yield) and Yi (the first effective quantum yield obtained from a rapid light curve) were observed, P. damicornis showed no visual signs of bleaching on the Imaging-PAM after this time. In A. nobilis and C. serailia, visual signs of bleaching over the 8 h period were accompanied by marked changes in F (light-adapted fluorescence yield), NPQ (non-photochemical quenching) and E _k (minimum saturating irradiance), as well as _PSII and Yi. These changes were most marked over the first 5 h. The most sensitive species was A. nobilis, which after 8 h at 33°C had reached a _PSII value of almost zero across its whole surface. Differential bleaching responses between polyps and coenosarc tissue were found in P. damicornis, but not in A. nobilis and C. serailia. NPQ increased with exposure time to 33°C in both the latter species, accompanied by a decreasing E _k, suggesting that the xanthophyll cycle is entrained as a mechanism for reducing the effects of the bleaching conditions.Communicated by L. Hagerman, Helsingør     

Use of pulse amplitude modulated (PAM) fluorometry for in situ measurements of photosynthesis in two Red Sea faviid corals

Measurements of the photosynthetic activity of symbiotic zooxanthellae in corals under natural growth conditions has been limited until recently, and this is one of the first reports on utilising a newly developed underwater pulse amplitude modulated (PAM) fluorometer (the Diving-PAM, Walz Gmbh, Germany) for such studies in situ. Photosynthetic responses to irradiance (photosynthetic photon flux, PPF) of the two faviid corals Favia favus (Forskål) and Platygyra lamellina (Ehrenberg) were measured while snorkelling or SCUBA diving (in August 1997), and we report here the results in terms of effective quantum yields of photosystem II (Y?) and estimated rates of photosynthetic electron transport (ETR, calculated as Y?×?0.5?× PPF?×?FA, where FA is the estimated fraction of light absorbed by the photosymbiont-containing tissue). Both species showed a reduction in Y with increasing actinic irradiances produced by the instrument above 500?μmol photons m^?2 s^?1, and the corresponding ETR values yielded apparently typical photosynthesis versus irradiance (P-I?) curves, which saturated between 1500 and 2000?μmol photons m^?2 s^?1. It was found that 30?s irradiation at each PPF level was sufficient to give optimal ETR values and, therefore, each P-I curve could be obtained within a few minutes. In situ point measurements from various areas of colonies under ambient light showed average ETR values within the range expected from the P-I curves. In order to test the Diving-PAM in an eco-physiologically relevant experiment, photosynthetic ETR versus PPF was measured for three sections of a large P. lamellina, each section of which received different natural irradiance levels. The results clearly demonstrated adaptations to the ambient light field in that vertical and downward-facing portions of the colony showed gradually lower maximal ETRs, steeper initial slopes of the P-I curves and, accordingly, lower light saturation points than upward-facing areas receiving higher light levels. Based on these trials, some evaluations are given as to the applicability of the Diving-PAM for photosynthetic measurements when monitoring similar corals.     

The relative contribution of dinoflagellate photosynthesis and stored lipids to the survivorship of symbiotic larvae of the reef-building corals

The long-distance dispersal of larvae provides important linkages between populations of reef-building corals and is a critical part of coral biology. Some coral planulae have symbiotic dinoflagellates (Symbiodinium spp.) that probably provide energy in addition to the lipids provisioned within the egg. However, our understanding of the influence of symbionts on the energy metabolism and survivorship of planulae remains limited. This study examines the relative roles of symbiotic dinoflagellate photosynthesis and stored lipid content in the survivorship of the developing stages of the corals Pocillopora damicornis and Montipora digitata. We found that survivorship decreased under dark conditions (i.e. no photosynthetic activity) for P. damicornis and M. digitata at 31 and 22 days after release/spawning, respectively. The lipid content of P. damicornis and M. digitata planulae showed a significant decrease, at a higher rate, under dark conditions, when compared with light conditions. When converted to energy equivalents, the available energy provided by the depletion of lipids could account for 41.9 and 84.7% of larval metabolism for P. damicornis (by day 31) and 38.4 and 90.1% for M. digitata (by day 21) under light and dark conditions, respectively. This finding indicates that not all energy requirements of the larvae are met by lipids: energy is also sourced from the photosynthetic activities of the symbiotic dinoflagellates within these larvae, especially under light conditions. In addition, the amounts of three main lipid classes (wax esters, triglycerides, and phospholipids) decreased throughout the experiment in the planulae of both species, with the wax ester content decreasing more rapidly under dark conditions than under light conditions. The observations that the planulae of both species derive considerable amounts of energy from wax esters, and that symbiotic dinoflagellates enable larvae to use their stores at lower rates, suggested that symbiotic dinoflagellates have the potential to extend larval life under light conditions.     

Patterns of Oil-Sediment rejection in corals

The patterns of oil-sediment rejection of 19 Caribbean hermatypic corals are identical to theri patterns of rejection of clean sediments. The rejection pattern is typical for coral species, and displays maximum and minimum rates dependent on the size and density of the oil-sediment particles. The viscosity of the oil determines the size of the oil-sediment particles. A coral's efficiency of rejection of sediment depends on the size and amount of the sediment particles. Oil drops 0.06 mm are removed by the coral's tissues. Physical contact with oil-sediment particles appears to be less harmful to corals than the toxic effects of oils.     

Turnover of photosynthetically fixed carbon in reef corals

Siderastrea siderea and Montastrea annularis were labeled in situ with NaH¹⁴CO₃. The corals were sampled over a period of 11 days and the radioactivity remaining in the ethanolsoluble and ammonia-soluble fractions measured. Total radioactivity in the corals fell to about 1/3 after one night and then to about 1/3 in the next 10 days. The ethanol-soluble radioactivity is probably converted to the less soluble, ammonia-extractable, material in the dark.Contribution No. 1514 from the University of Miami, Rosenstiel School of Marine and Atmospheric Science, 10 Rickenbacker Causeway, Miami, Florida 33149, USA.     

Age-size plasticity for reproduction in monocarpic plants

Empirical and theoretical investigations of monocarpy have usually addressed the question of minimum or threshold sizes for reproduction. However, the range of flowering sizes observed in many monocarpic species is extraordinarily large (well beyond what can be called a "threshold"), and the sizes of flowering and nonflowering plants may overlap greatly. We attempt to explain these reproductive patterns in terms of optimal reaction norms predicted by simple deterministic life history models. We assume that individuals differ in their growth trajectories due to the heterogeneous quality of microsites and ask how the optimal age and size at flowering varies with environmental variation in growth and for different assumptions about fecundity and mortality. Under two very different growth functions (one with no age- or size-related decline in growth rate and another with such a decline as size approaches an asymptote), the optimal reaction norms imply considerable plasticity for size at reproduction, particularly when poor growth is associated with higher mortality or lower asymptotic size. Deterministic models such as these may be more applicable to long-lived than to short-lived monocarps, because fitness potential should be less affected by stochastic variability in yearly growing condition in the former than in the latter. We consider the case of a tropical monocarpic and masting tree species, Cerberiopsis candelabra (Apocynaceae), and show that our model results can account for wide ranges of reproductive size and overlap in size of flowering and nonflowering plants, in accord with observation. We suggest that empirical attention to norms of reaction across growth environments will be a more profitable approach than investigation of size thresholds per se.     

Experimental tests of suspension feeding in Atlantic reef corals

The ability of 15 species of Atlantic reef corals to act as suspension feeders was demonstrated by their removal of suspended particles from sea water in culture vessels. Mean clearance rates varied from 16.6 to 145.5 ml water cleared/h/cm² of live coral tissue. The lowest rates was found in Porites porites which is primarily a tentacle feeder, and the highest in Diploria clivosa which acts as both a tentacle feeder and suspension feeder. Rates of particle clearance in Agaricia agaricites, which is primarily a suspension feeder, were influenced by current velocity and type of food.     

Sexual and asexual production of planulae in reef corals

Electrophoresis was used to provide genetic evidence of the mode of production of brooded planulae in each of four species of scleractinian coral collected from the central region of the Great Barrier Reef during September, October and November 1984. Comparisons were made of the multi-locus genotypes of planulae and their broodparents for two ahermatypic (non zooxanthellate) species,Tubastraea diaphana andT. coccinea and two hermatypic (zooxanthellate) species,Acropora palifera andSeriatopora hystrix. For both ahermatypic species, all planulae were found to be genetically identical to their broodparents, including 91 planulae which were heterozygous for at least one locus. These results are consistent with asexual (ameiotic) reproduction. In contrast, non-parental genotypes were detected in the majority of hermatypic broods, which is consistent with expectations for sexual reproduction with at least some outcrossing. These data confirmed that brooded planulae may be produced both sexually and asexually and countered the suggestion that electrophoretic studies of hermatypic corals may be weakened by the contaminating effect of the enzymes of their symbiotic xooxanthellae.Contribution No. 306 of the Australian Institute of Marine Science     

Measuring mucus thickness in reef corals using a technique devised for vertebrate applications

A method previously used to measure thickness of the surface mucus layer (SML) of the mammalian gastrointestinal tract has been applied to the SML of reef corals. It involves manual measurement of mucus thickness using a micromanipulator and fine glass needle (micropipette) and is non-destructive to the coral, meaning that repeated measurements can be taken. A measurable mucus layer was recorded in all cases in the study, which comprised 450 individual thickness measurements from four coral species. Mucus thickness ranged from 145 to 700 μm. Thus, whatever dynamic processes control mucus synthesis, secretion to the tissue surface and subsequent release into the water column, a continuous mucosal barrier is maintained. A change in SML thickness was recorded as a response to aerial exposure during the natural tidal cycle and to solar exposure-induced bleaching, although the response due to bleaching varied between two studied species. The technique is rapid, cost-effective and a simple means of assessing coral SML thickness, a variable that shows significant variation in relation to environmental conditions and is likely to be an important health indicator in these organisms.     

Allometric variability of heavy metals in corals of Lakshadweep islands

Heavy metal contents of the corals collected from the Lakshadweep archipelago are discussed in the light of allometric variations. Plots of log flesh weight versus log flesh metal concentration exhibited negative correlations for all the nine metals studied. Metals with rapid exchange rate are likely to show such negative correlation and in such situations the elimination rate far exceeds the metal accumulation rate. Similar plots in the skeletal component showed positive correlations. The only linear relationships were observed in log flesh density versus log partition coefficient plots. Comparative flesh enrichment of metals were found to decrease against increasing flesh weights (for both essential and non-essential metals), indicating a regulation of flesh metal burden beyond certain limits. In the skeletal component, anthropogenic input as well as competitive interactions between metal ions having radii similar to that of Ca²⁺ paves way for metal adsorption in calcite/aragonite skeletal lattice of corals.     

Diel fluctuations of mycosporine-like amino acids in shallow-water scleractinian corals

Reports of bathymetric decrease in the total mycosporine-like amino acid (MAA) concentration of benthic invertebrates suggest that light gradients may be important determinants of MAA content. With the pronounced diel light changes, distinct temporal variations in MAA concentrations might also be expected. We examined the changes in the abundance of MAA in three shallow-water scleractinian corals, Pavona divaricata, Galaxea fascicularis and Montipora digitata from Okinawa, Japan, in relation to daily cycles in solar radiation and tested whether the species have different capabilities for protection against UVR depending on their MAA composition. The results show that symbiotic algae freshly isolated from the investigated coral species do not contain MAAs and that distribution of these compounds resided only within the animal tissue. Total MAA content in the tissue of P. divaricata, G. fascicularis and M. digitata rose rapidly at midday and significantly dropped at night. The observed variations were by a factor of two and, thus, very dramatic. For all the investigated coral species, total MAA concentrations were significantly correlated with the diurnal cycle in solar radiation, during both winter and summer seasons. Seawater temperature was significantly correlated with MAA levels only in the June experiment, but represented no more than 20% of the MAA variation in all three coral species, whereas solar radiation explained 60–70% of the MAA fluctuations. This suggests that MAAs are an integral component of the hard corals biochemical defense system against high solar irradiance stress. The diurnal increase in total MAA concentrations was due to an increase in the concentration of imino-MAA species of up to 2–2.5-fold of their pre-dawn values. In contrast, the oxocarbonyl-MAA mycosporine-glycine (Myc-Gly) showed the lowest (Tukey–Kramer HSD test: P<0.05) values at midday, compared to afternoon and night hours. Analysis of diel changes in chlorophyll fluorescence and chlorophyll a content of the investigated species revealed that P. divaricata and G. fascicularis were less sensitive to the high levels of ambient irradiance compared to M. digitata. In P. divaricata and G. fascicularis, Myc-Gly, an MAA with an antioxidant function, is the most abundant MAA, contributing about 70% to the total MAA pool, whereas the major MAA factions in M. digitata were represented by oxidatively robust imino-MAAs. We speculate that MAAs furnish scleractinian corals with protection from biologically damaging ultraviolet radiation through both the direct sunscreening activity of imino-MAAs and the antioxidant properties of oxocarbonyl-MAAs and suggest that the predominance, in the host tissue, of MAA species with an antioxidant ability may render corals more tolerant to high photosynthetically active and ultraviolet radiation.Communicated by T. Ikeda, Hakodate     

Photoadaptation of photosynthesis in Gonyaulax polyedra

Gonyaulax polyedra Stein exhibited a combination of photoadaptive strategies of photosynthesis when only a single environmental variable, the light intensity during growth, was altered. Which of several biochemical/physiological adjustments to the light environment were employed depended on the level of growth irradiance. The photoadaptive strategies employed over any small range of light levels appeared to be those best suited for optimizing photosynthetic performance and not photosynthetic capacity. (Photosynthetic performance, P _i, is defined as the rate of photosynthesis occurring at the level of growth irradiance.) Among all photosynthetic parameters examined, only photosynthetic performance showed a consistent correspondence to growth rates of G. polyedra. Above 3500 to 4000 W cm^-2, where photosynthetic performance was equal to photosynthetic capacity, cells were not considered light-limited in either photosynthesis or growth. At these higher light levels, photosynthetic perfomance, cell volume, growth rates and respiration rates remained maximal; photosynthetic pigment content varied only slightly, while the photosynthetic capacity of the cells declined. At intermediate light levels (3000 to 1500 W cm^-2), photosynthesis, not growth, was light-limited, and photoadaptive strategies were induced which enhance absorption capabilities and energy transfer efficiencies of chlorophyll a to the reaction centers of G. polyedra. Photosynthetic capacity remained constant at about 280 mol O₂ cm^-3 h^-1, while photosynthetic performance ranged from 100 to 130 mol O₂ cm^-3 h^-1. Major increases in photosynthetic pigments, especially peridinin-chlorophyll a-proteins and an unidentified chlorophyll c component, accompanied photoadaptation to low irradiances. Maximal growth rates of 0.3 divisions day^-1 were maintained, as were respiration rates of about-80 mol O₂ cm^-3 h^-1 and cell volumes of about 5.4×10^-8 cm^-3 cell^-1. Below about 1250 W cm^-2, photosynthesis in G. polyedra was so light-limited that photosynthetic performance was unable to support maximal growth rates. Under these conditions, G. polyedra displayed photostress responses rather than photoadaptive strategies. Photostress was manifested as reduced cell volumes, slower growth, and drastic reductions in pigmentation, photosynthetic capacity, and rates of dark respiration.     

Depth analysis of fatty acids in two caribbean reef corals

Total fatty acid compositions of colonies of two hermatypic, reef-building corals collected during the day-time over a depth range of 21 m were determined to assess the effect of depth-related environmental factors upon the lipid content of these organisms. No systematic changes were found, suggesting a steady-state balance between algal and animal lipogenesis in these symbiotic partnerships. Stephanocoenia michelinii, a day and night feeder, contained lipids indicative of external dietary sources such as copepods, whereas Montastrea annularis, a night feeder, did not.     

Reproductive patterns of scleractinian corals in the northern Red Sea

The majority of published accounts on scleractinian coral reproduction are from the tropical Pacific and Caribbean, with very little information known about Red Sea species. This report examines variation in reproductive mode in 24 species of hermatypic corals (belonging to seven families) in the Gulf of Eilat, Red Sea. Eighteen species are hermaphroditic broadcasters, two are hermaphroditic brooders and three are gonochoric broadcasters. In the Pocilloporidae, the gonads project into the body cavity, while in the other six families the gonads reside inside the mesenteries. The number of gonads per polyp in broadcasting species follows family or genus lines. Fecundity (eggs per polyp) increases with polyp size. Brooding species usually exhibit one or two gonads per polyp and each gonad contains only one to three oocytes. Oocyte size varies widely and does not relate to mode of reproduction. The largest oocytes (diameter = 450 μm) occur in the brooding coral Alveopora daedalea and in broadcasting species of the genus Acropora (diameter = 420 μm). Gonad morphology and gonochorism versus hermaphroditism appear to be constrained phylogenetically at the family or genus level. Lastly, this report compares the data presented for Red Sea scleractinian species with the data available on scleractinian corals from other geographical regions. Received: 2 February 1993 / Accepted: 9 March 1998     

Dynamics of a coral reef community after mass mortality of branching <Emphasis Type="Italic">Acropora</Emphasis> corals and an outbreak of anemones

Dynamics of a coral reef community at Tiao-Shi Reef, southern Taiwan were studied using permanent transects to examine coral recovery and successive cascades to collapse stage resulting from chronic anthropogenic impacts and typhoons. Three distinct zones were recognized within a relatively small study area (250 m across) formerly dominated by large stands of branching Acropora corals. The first zone still retains the dominance of branching Acropora corals, although they show a significant decreasing tendency. The second zone exhibits recovery with a significant increase in branching Montipora stellata, which is recruited and grows faster than branching Acropora corals. The third zone is occupied by anemone, Condylactis sp., and demonstrates a stable phase of coral deterioration without recovery. Such differences in coral reef community dynamics within a small spatial scale illustrate mosaic dynamics which have resulted from degradation of the water quality, patchy mortality of large branching Acropora thickets caused by typhoons, the rapid asexual fragmentation and growth of M. stellata making it a successful colonizer, and occupation by anemone, Condylactis sp., together with unstable remnants of dead Acropora rubbles have not allowed coral recruits to survive.