Visibly healthy corals exhibit variable pigment concentrations and symbiont phenotypes

Understanding the natural variability of photosynthetic pigment ranges and distributions in healthy corals is central to evaluating how useful these measurements are for assessing the health and bleaching status of endosymbiotic reef-building corals. This study examined the photosynthetic pigment variability in visibly healthy Porites lobata and Porites lutea corals from Kaneohe Bay, Hawaii and explored whether pigment variability was related to the genetic identity or phenotypic characteristics of the symbionts. Concentrations of the photosynthetic pigments chlorophyll a, peridinin, chlorophyll c ₂ , diadinoxanthin, diatoxanthin, β,β-carotene and dinoxanthin were quantified using high-performance liquid chromatography (HPLC). Pigment concentrations were found to range 1.5–10 fold in colonies of each species at similar depths (0–2, 2–4, 10–15 and 19–21 m). Despite the high pigment variability, pigment ratios for each species were relatively conserved over the 0–21 m depth gradient. The genetic identity of the symbiont communities was examined for each colony using 18S nuclear ribosomal DNA (nrDNA) restriction fragment length polymorphisms. All colonies contained symbionts belonging to clade C. The density and phenotypic characteristics of the symbionts were explored using flow cytometry, and fluorescence and side scatter (cell size) properties revealed phenotypically distinct symbiont subpopulations in every colony. The symbiont subpopulations displayed pigment trends that may be driven by acclimatization to irradiance microenvironments within the host. These results highlight the biological complexity of healthy coral–symbiont associations and the need for future research on pigments and symbiont subpopulation dynamics.     

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.     

PHOTOSYNTHESIS IN SOME ALCYONACEAN CORALS

Burkholder , Paul R., and Lillian M. Burkholder . (Brooklyn Botanic Garden, Brooklyn, New York.) Photosynthesis in some Alcyonacean corals. Amer. Jour. Bot. 47(10) : 866–872. Illus. 1960.—Quantitative production of oxygen in bottles, and amounts of chlorophyll and carotenoid pigments were studied in selected species of Alcyonacean corals, the stony coral Porites, porites, and turtle grass, Thalassia testudinum, in Puerto Rico. Photosynthetic production exceeds the consumption of oxygen in combined respiration of the corals and their contained algal zooxanthellae, both in short-term light experiments and over 24 hr. of natural, alternating day and night. Observations of oxygen production in 8 species of corals, exposed to full sunlight in 1 ft. of water, gave assimilation values of 0.9 mg. O₂ (equivalent to 0.27 mg. C fixed) per mg. chlorophyll a per hour. A maximum assimilation value of 1.56 was observed in Antillogorgia turgida. From data on the increased oxygen it was calculated that from 0.33 to 1.65 mg. residual fixed carbon per g. of fresh coral were accumulated in Porites porites and Antillogorgia turgida during a day and night period of 24 hr. Photosynthesis of turtle grass in the same environment was more than 3 times the rate of these corals. The zooxanthellae contain carotenoids in abundance, along with chlorophylls a and c. On a unit of chlorophyll a basis, the photosynthetic systems of corals appear to be less active in reduction of carbon dioxide in bottles than some other photosynthetic systems of the sea, such as turtle grass and phytoplankton observed under optimum conditions.     

Effects of iron on growth and reflectance spectrum of the bloom‐forming cyanobacterium Microcystis viridis

Iron is an important factor in algal blooms because it is involved in cyanobacterial pigment biosynthesis and therefore has the ability to influence the pigment status of algal cells. This role in pigment biosynthesis offers the opportunity for rapid monitoring of iron availability to cyanobacteria through spectral reflectance characterization. In the present study, the freshwater cyanobacterium Microcystis viridis was cultured with different levels of iron. Cell density, cellular content of iron and photosynthetic pigments, and spectral reflectivity of M. viridis were determined daily during the course of the culture experiment. The results showed that at the lowest iron concentration (0.01 μM) the growth of M. viridis was seriously limited, and the maximal cell density was only approximately 6.4% of the density observed with an iron concentration of 18 μM. Iron availability dramatically affected chlorophyll a, carotenoid and phycocyanin content, with the greatest impact on chlorophyll a. The iron‐induced changes in content and ratios of pigments were detectable through spectral reflectance. Eleven spectral indices previously developed for the estimation of concentrations and/or ratios of pigments and a newly proposed chlorophyll a/phycocyanin index were found to be suitable for generating sensitive regression models between cellular iron content and spectral parameters. The comprehensive application of key sensitive spectral indices and regression equations should help to support monitoring and diagnosis of iron availability to cyanobacteria via remote sensing.     

Corals as light collectors: an integrating sphere approach

An integrating sphere was used to estimate the fraction of the incident quantum flux absorbed by a coral colony placed within it. This method allows one to examine the in vivo light absorption of intact coral colonies. We used this method to study effects of colony morphology, size, and photoacclimation status on the light harvesting efficiency by the zooxanthellae. Light absorption per unit of coral surface area decreased with increase in colony size, with a clear effect of different coral morphologies. In branched colonies, shading among branches reduced the absorbed light per unit area and per zooxanthellae. Photoacclimation to low light resulted in increased cellular chlorophyll concentrations in the zooxanthellae. In shade acclimated colonies, areal chlorophyll concentrations increased significantly, leading to more overlap among the optical cross-sections of pigments within cells and mutual shading among cells. These package effects showed up as a decrease in the in vivo, chlorophyll-a specific, spectral average, effective optical cross-section, a*. An integrating sphere is a useful tool for collecting optical information on corals.     

DETECTION OF CHLOROPHYLLS c1, c2 AND c3 IN PIGMENT EXTRACTS OF PRYMNESIUM PARWM (PRYMNESIOPHYCEAE)1

Three chlorophyll c-type pigments were separated by reversed-phase high Performance liquid chromatography and thin-layer chromatography from pigment extracts of the prymnesiophyte, Prymnesium parvum Carter. Based on spectral characteristics, retention times, and comparison with reference pigments isolated from the diatom Phaeodactylum tricornutum Bohlin, two of these pigments were identijied as chlorophyll c₁ and c₂. The other pigment was identified by its absorption spectrum and thin-layer chromatography retention times as the newly described chlorophyll c₃. However, in other prymnesiophytes so far examined, chlorophyll c₁ and chlorophyll c₃ were present with no chlorophyll c_l. The discovery of chlorophyll c₃ with chlorophyll c₁ and chlorophyll c₃ in Prymnesium parvum therefore represents the first report of this combination of pigments in prymnesiophytes.     

Optical spectra and pigmentation of Caribbean reef corals and macroalgae

 Coastal reef degradation and widespread bleaching of corals, i.e. loss of pigments and/or symbiotic zooxanthellae, is increasing globally. Remote sensing from boats, aircraft or satellites has great potential for assessing the extent of reef change, but will require ground-verified spectral algorithims characteristic of healthy and degraded reef populations. We collected seven species of Caribbean reef corals and also representative macroalgae from reefs near Lee Stocking Island, Bahamas and quantified their pigments using high performance liquid chromatography. We also measured the fluorescence and reflectance spectra of corals and macroalgae using an in situ benthic spectrofluorometer. In visibly pigmented (unbleached) coral from 4 to 5 m depth, the mean (±SD) surface density of pigments (3.0±1.3 μg chlorophyll-a cm^-2 and 2.1±0.7 μg peridinin cm^-2) was similar between colonies of the same species, but differed among species. The mean quantity of pigment per zooxanthella (1.8±0.9 pg chl-a cell^-1 and 1.4±0.7 pg peridinin cell^-1) also differed among species and sometimes between colonies of the same species. Chl-a and peridinin densities per surface area of coral were positively correlated. When excited with blue light (480 nm), macroalgae and corals had typical chlorophyll fluorescence with a peak at 680 nm and a smaller shoulder peak at 730 to 740 nm. Most corals, unlike macroalgae, also had distinct fluorescence peaks between 500 and 530 nm. In visibly bleached corals 680 nm fluorescence was greatly reduced in amplitude. Pigmented coral, under natural lighting conditions, had a reflected light peak at about 570 nm. Reflectance increased over all wavelengths in bleached corals, with the greatest increase at the wavelengths where chlorophyll and accessory pigments absorb light, i.e. 670 and 450 to 550 nm. Both fluorescence and reflectance spectra appear promising to remotely differentiate between pigmented and bleached coral and between coral and macroalgae. Accepted: 15 March 1999     

Spectral reflectance indices and pigment functions during leaf ontogenesis in six subtropical landscape plants

Pigment combinations are regulated during leaf ontogenesis. To better understand pigment function, alterations in chlorophyll, carotenoid and anthocyanin concentrations were investigated during different leaf development stages in six subtropical landscape plants, namely Ixora chinensis Lam, Camellia japonica Linn, Eugenia oleina Wight, Mangifera indica L., Osmanthus fragrans Lowr and Saraca dives Pierre. High concentrations of anthocyanin were associated with reduced chlorophyll in juvenile leaves. As leaves developed, the photosynthetic pigments (chlorophyll and carotenoid) of all six species increased while anthocyanin concentration declined. Chlorophyll fluorescence imaging of Φ_PSII (effective quantum yield of PSII) and of NPQ (non-photochemical fluorescence quenching) and determination of electron transport rate-rapid light curve (RLC) showed that maximum ETR (leaf electron transport rate), Φ_PSII and the saturation point in RLC increased during leaf development but declined as they aged. Juvenile leaves displayed higher values of NPQ and Car/Chl ratios than leaves at other developmental stages. Leaf reflectance spectra (400–800 nm) were measured to provide an in vivo non-destructive assessment of pigments in leaves during ontogenesis. Four reflectance indices, related to pigment characters, were compared with data obtained quantitatively from biochemical analysis. The results showed that the ARI (anthocyanin reflectance index) was linearly correlated to anthocyanin concentration in juvenile leaves, while a positive correlation of Chl NDI (chlorophyll normalized difference vegetation index) to chlorophyll a concentration was species dependent. Photosynthetic reflectance index was not closely related to Car/Chl ratio, while a structural-independent pigment index was not greatly altered by leaf development or species. Accordingly, it is suggested that the high concentration of anthocyanin, higher NPQ and Car/Chl ratio in juvenile leaves are important functional responses to cope with high radiation when the photosynthetic apparatus is not fully developed. Another two leaf reflectance indices, ARI and Chl NDI, are valuable for in vivo pigment evaluation during leaf development.     

Host pigments: potential facilitators of photosynthesis in coral symbioses

Reef‐building corals occur as a range of colour morphs because of varying types and concentrations of pigments within the host tissues, but little is known about their physiological or ecological significance. Here, we examined whether specific host pigments act as an alternative mechanism for photoacclimation in the coral holobiont. We used the coral Montipora monasteriata (Forskål 1775) as a case study because it occurs in multiple colour morphs (tan, blue, brown, green and red) within varying light‐habitat distributions. We demonstrated that two of the non‐fluorescent host pigments are responsive to changes in external irradiance, with some host pigments up‐regulating in response to elevated irradiance. This appeared to facilitate the retention of antennal chlorophyll by endosymbionts and hence, photosynthetic capacity. Specifically, net P_max Chl a^?1 correlated strongly with the concentration of an orange‐absorbing non‐fluorescent pigment (CP‐580). This had major implications for the energetics of bleached blue‐pigmented (CP‐580) colonies that maintained net P_max cm^?2 by increasing P_max Chl a^?1. The data suggested that blue morphs can bleach, decreasing their symbiont populations by an order of magnitude without compromising symbiont or coral health.     

Spectral discrimination of coral reef benthic communities

Effective identification and mapping of coral reef benthic communities using high-spatial and -spectral resolution digital imaging spectrometry requires that the different communities are distinguishable by their spectral reflectance characteristics. In Kaneohe Bay, Oahu, Hawaii, USA, we collected in situ a total of 247 spectral reflectances of three coral species (Montipora capitata, Porites compressa, Porites lobata), five algal species (Dictyosphaeria cavernosa, Gracilaria salicornia, Halimeda sp., Porolithon sp., Sargassum echinocarpum) and three sand benthic communities (fine-grained carbonate sand, sand mixed with coral rubble, coral rubble). Major reflectance features were identified by peaks in fourth derivative reflectance spectra of coral (at 573, 604, 652, 675 nm), algae (at 556, 601, 649 nm) and sand (at 416, 448, 585, 652, 696 nm). Stepwise wavelength selection and linear discriminant function analysis revealed that spectral separation of the communities is possible with as few as four non-contiguous wavebands. These linear discriminant functions were applied to an airborne hyperspectral image of a patch reef in Kaneohe Bay. The results demonstrate the ability of spectral reflectance characteristics, determined in situ, to discriminate the three basic benthic community types: coral, algae and sand. Accepted: 12 January 2000     

冠层水平互花米草叶片光合色素含量的高光谱遥感估算模型

以闽江河口鳝鱼滩湿地互花米草(Spartina alterniflora)的实测冠层高光谱反射率和叶片光合色素含量(LPPC)为数据源,在分析LPPC与原始光谱反射率、一阶导数光谱反射率、22种已报道光谱指数和14种新构建的植被指数相关性的基础上,利用直线回归、指数回归、对数回归以及乘幂回归方法,系统地比较了36种植被指数在估算互花米草LPPC中的表现。研究表明:(1)一阶导数光谱反射率组合的植被指数用于估算互花米草的LPPC优于原始光谱反射率;(2)红边区域一阶导数光谱是估测互花米草LPPC的最佳波段;(3)对于单一色素含量的估算,叶绿素a(Chla)的最佳估算指数为FDNDVI[723,703];叶绿素b(Chlb)的最佳估算指数为FDRVI[723,525];类胡萝卜素(Cars)的最佳估算指数为FDNDVI[723,703];(4)对于使用统一参量同时估算Chla、Chlb、Cars,由FDRVI[723,703]建立的对数估算模型效果最佳。研究成果可为湿地植物生化参量反演提供参考,也可为闽江河口湿地入侵种互花米草的动态监测和生态评估管理提供有力的科学依据。     

Surface reflectance properties of Antarctic moss and their relationship to plant species, pigment composition and photosynthetic function

In this study the variations in surface reflectance properties and pigment concentrations of Antarctic moss over species, sites, microtopography and with water content were investigated. It was found that species had significantly different surface reflectance properties, particularly in the region of the red edge (approximately 700 nm), but this did not correlate strongly with pigment concentrations. Surface reflectance of moss also varied in the visible region and in the characteristics of the red edge over different sites. Reflectance parameters, such as the photochemical reflectance index (PRI) and cold hard band were useful discriminators of site, microtopographic position and water content. The PRI was correlated both with the concentrations of active xanthophyll‐cycle pigments and the photosynthetic light use efficiency, F_v/F_m, measured using chlorophyll fluorescence. Water content of moss strongly influenced the amplitude and position of the red‐edge as well as the PRI, and may be responsible for observed differences in reflectance properties for different species and sites. All moss showed sustained high levels of photoprotective xanthophyll pigments, especially at exposed sites, indicating moss is experiencing continual high levels of photochemical stress.     

OCCURRENCE OF PHYIWATED CHLOROPHYLL c IN ISOCHRYSIS GALBANA AND ISOCHRYSIS SP. (CLONE T-ISO) (PRYMNESIOPHYCEAE)1

The pigment composition of two clones of Isochrysis galbana Parke (CCMP 1323 and CCAP 927/1), and Isochrysis sp. (clone T-ISO) was analyzed by high-performance liquid chromatography using a polymeric octadecylsilica column. Fluorescent peaks with retention times higher than chlorophyll a were detected for all three clones. The corresponding pigments were isolated and characterized in terms of their visible absorbance and fluorescence spectra. The pigments were similar to phytol-substituted chlorophyll c, previously isolated from Emiliania huxleyi (Lohm.) Hay and Mohler and other species containing chlmophyll c₃. The presence of phytol-substituted chlorophyll c in I. galbana which lacked chlorophyll c₃, increases the diversity of chlorophyll patterns for the Haptophyta, which can be grouped, at present, into six different pigment types. This is the jrst observation of a haptophyte containing the apolar phytylated chlorophyll c-like pigment but lacking chlorophyll c₃.     

PHOTOSYNTHETIC PIGMENTS IN FIFTY-ONE SPECIES OF MARINE DIATOMS1

The photosynthetic pigments of 51 species (71 isolates) of tropical and sub-tropical diatoms from 13 out of 22 families were examined. These were the Thalassiosiraceae, Melosiraceae, Coscinodiscaceae, Rhizosoleniaceae, Biddulphiaceae, Chaetoceraceae, Lithodesmiaceae, Eupodiscaceae, Cymatosiraceae, Diatomaceae, Naviculaceae, Nitzschiaceae and Phaeodactylinaceae. Pigments were analyzed by cellulose and polyethylene thin-layer chromatography (TLC) and reverse-phase high-performance thin-layer chromatography (HPTLC). All species contained chlorophylls a and c₂ and the carotenoids carotene, fucoxanthin, diatoxanthin and diadinoxanthin. In addition, 14 species (20 isolates) contained one or more of four minor carotenoids, which were not identified further. One species, Thalassiothrix heteromorpha, contained small amounts of a 19′-butanoyloxyfucoxanthin-like pigment, in addition to fucoxanthin. Chlorophyll c₂ was present in all the diatoms tested and occurred together with chlorophyll c₁ in 88% of them. The presence of both chlorophylls c₁ and c₂ therefore can no longer be considered a universal characteristic of the diatom class. Where chlorophyll c₁ was absent or occurred in trace amounts only (8 species, 11 isolates), it was usually replaced by a new chlorophyll c pigment designated chlorophyll c₃, recently characterized from several prymnesiophytes and one chrysophyte. Exceptions were Nitzschia closterium (CS-114), which contained only chlorophyll c₂, and Nitzschia bilobata (CS-47), which contained all three chlorophylls (c₁, c₂ and c₃) in approximately equal amounts. Five species that contained chlorophylls c₁ and c₂ also contained chlorophyll c₃ in trace quantities Quantitative pigment analyses of the 71 isolates showed that chlorophyll concentrations ranged from 0.02 μg. 10⁶ cells^?1 in the smallest diatom, Extubocellulus spinifer, to 174.4 μg. 10⁶ cells^?1 in one of the largest diatoms, Coscinodiscus sp. under the standard growth conditions used. The mean molar ratio of chlorophyll a:c in the 72 isolates was 3.33, with a range of 1.65–7.25. The close similarity between diatom and prymnesiophyte pigmentation was confirmed. Each class has three patterns of pigmentation: viz species with chlorophylls c₁ and c₂ and‘true’fucoxanthin, species with chlorophylls c₃ and c₂ and‘true’fucoxanthin, and species with chlorophylls c₃ and c₂ and fucoxanthin derivatives.     

The effects of elevated temperature on the photosynthetic efficiency of zooxanthellae in hospite from four different species of reef coral: a novel approach

Bleaching of reef corals is a phenomenon linked to temperature stress which involves loss of the symbiotic algae of the coral, which are known as zooxanthellae, and/or loss of algal pigments. The photosynthetic efficiency of zooxanthellae within the corals Montastrea annularis, Agaricia lamarki, Agaricia agaricites and Siderastrea radians was examined by pulse-amplitude modulation fluorometry (PAM) during exposure to elevated temperatures (30–36°C). Zooxanthellae within M. annularis and A. lamarki were found to be more sensitive to elevated temperature, virtually complete disruption of photosynthesis being noted during exposure to temperatures of 32 and 34°C. The photosynthetic efficiency of zooxanthellae within S. radians and A. agaricites decreased to a lesser extent. Differences in the loss of algal cells on an aerial basis and in the cellular chlorophyll concentration were also found between these species. By combining the non-invasive PAM technique with whole-cell fluorescence of freshly isolated zooxanthellae, we have identified fundamental differences in the physiology of the symbionts within different species of coral. Zooxanthellae within M. annularis appear to be more susceptible to heat-induced damage at or near the reaction centre of Photosystem II, while zooxanthellae living in S. radians remain capable of dissipating excess excitation energy through non-photochemical pathways, thereby protecting the photosystem from damage during heat exposure.     

A new form of chlorophyll C involved in light-harvesting

A new form of chlorophyll c has been isolated from the pyrmnesiophyte Pavlova gyrans Butcher. This pigment is spectrally similar to chlorophyll c₂, but all the absorption maxima (454, 583, and 630 nm in diethyl ether) are shifted 4 to 6 nanometers to longer wavelengths. The new pigment can be separated from other chlorophyll c-type pigments by reversed-phase high performance liquid chromatography and thin layer chromatography. Both chlorophylls c₁ and c₂ are found with the new chlorophyll c pigment in P. gyrans, and it has also been detected in the chrysophyte Synura petersenii Korsh. The light-harvesting function of the new chlorophyll c pigment is indicated by its presence along with chlorophyll c₁ and c₂ in a light-harvesting pigment-protein complex isolated from P. gyrans in which chlorophyll c pigments efficiently transfer absorbed light energy to chlorophyll a.     

Quantification of total and particulate dimethylsulfoniopropionate (DMSP) in five Bermudian coral species across a depth gradient

The symbiotic dinoflagellate microalgae of corals (Symbiodinium spp.) contain high concentrations of dimethylsulfoniopropionate (DMSP), a multifunctional metabolite commonly found in many species of marine algae and dinoflagellates. A photoprotective antioxidant function for DMSP and its breakdown products has often been inferred in algae, but its role(s) in the coral–algal symbiosis remains elusive. To examine potential correlations between environmental and physiological parameters and DMSP, total DMSP (DMSP_t, from the host coral and zooxanthellae), particulate DMSP (DMSP_p, from the zooxanthellae only), coral surface area, and total protein, as well as zooxanthellae density, chlorophyll concentration, cell volume and genotype (i.e., clade) were measured in five coral species from the Diploria-Montastraea-Porites species complex in Bermuda along a depth gradient of 4, 12, 18, and 24 m. DMSP_t concentrations were consistently greater than DMSP_p concentrations in all species suggesting the possible translocation of DMSP from symbiont to host. D. labyrinthiformis was notably different from the other corals examined, showing DMSP_p and DMSP_t increases (per coral surface area or tissue biomass) with increasing water depth. However, overall, there were no consistent depth-related patterns in DMSP_p and DMSP_t concentrations. Further research, investigating dimethylsulfide (DMS), dimethylsulfoxide, and acrylate levels and DMSP-lyase activity in correlation with other biomarker endpoints that have been shown to be depth (i.e., temperature and light) responsive are needed to substantiate the significance of these findings.     

Recovery of the coral Montastrea annularis in the Florida Keys after the 1987 Caribbean “bleaching event”

Many reef-building corals and other cnidarians lost photosynthetic pigments and symbiotic algae (zooxanthellae) during the coral bleaching event in the Caribbean in 1987. The Florida Reef Tract included some of the first documented cases, with widespread bleaching of the massive coral Montastrea annularis beginning in late August. Phototransects at Carysfort Reef showed discoloration of >90% of colonies of this species in March 1988 compared to 0% in July 1986; however no mortality was observed between 1986 and 1988. Samples of corals collected in February and June 1988 had zooxanthellae densities ranging from 0.1 in the most lightly colored corals, to 1.6x10⁶ cells/cm² in the darker corals. Minimum densities increased to 0.5x10⁶ cells/cm² by August 1989. Chlorophyll-a content of zooxanthellae and zooxanthellar mitotic indices were significantly higher in corals with lower densities of zooxanthellae, suggesting that zooxanthellar at low densities may be more nutrientsufficient than those in unbleached corals. Ash-free dry weight of coral tissue was positively correlated with zooxanthellae density at all sample times and was significantly lower in June 1988 compared to August 1989. Proteins and lipids per cm² were significantly higher in August 1989 than in February or June, 1988. Although recovery of zooxanthellae density and coral pigmentation to normal levels may occur in less than one year, regrowth of tissue biomass and energy stores lost during the period of low symbiont densities may take significantly longer.     

Acclimation of the Hermatypic Coral Stylophora pistillata to Bright Light

Photoacclimation dynamics to bright light was studied in the symbiotic reef-building coral Stylophora pistillata. Coral colonies were collected from shallow shaded sites (2 m, 40–20% PAR_s) from a fringing reef at Sesoko Island, Okinawa, Japan. Outer branches were broken off from the colonies and placed in an outdoor aquarium until the start of the experiments. After maintenance of the branches in an aquarium under a light intensity of 30% PAR_s for 30 days (experiment 1) or for 90 days (experiment 2), the samples were exposed to 95% PAR_s for 120 days in the same aquarium. The population density of zooxanthellae, chlorophyll concentration, locations of zooxanthellae, proliferating zooxanthellae frequency (PZF), and degrading zooxanthellae frequency (DZF) were examined. It was shown that after acclimation of coral branches to bright light, the population density of zooxanthellae, chlorophyll concentration calculated per 1000 polyps, and chlorophyll concentration in zooxanthellae decreased. The size of zooxanthellae significantly decreased. A decrease in the population density of zooxanthellae was detected by the eighth day of acclimation, and stabilization in the density of the symbionts occurred in the period from the 40th to the 60th day of the experiment. The chlorophyll concentration in zooxanthellae significantly decreased by the second day of exposure to bright light and stabilized by the fourth day. The PZF level sharply dropped on the second day, while the DZF level sharply increased and was higher than the PZF level for 40 days of exposure to bright light. We conclude, therefore, that the population density of zooxanthellae is regulated by the rates of two processes: cell division and the cell degradation.     

Photo-acclimation of the hermatypic coral Stylophora pistillata while subjected to either starvation or food provisioning

This study investigated the photo-acclimation capacity of the coral Stylophora pistillata (Esper). Outer branches of coral colonies, taken from 2 m, were subjected to 90, 20, or 3% of incident surface photosynthetic active radiation (PAR(0)), or kept in total darkness. The corals were maintained either in filtered seawater (i.e., under starvation), or in seawater that had daily additions of zooplankton (rotifers). The experiments were maintained for 31 days. Zooxanthellae population densities and chlorophyll concentrations increased in S. pistillata fragments subjected to 20 and 3% PAR(0). The zooxanthellae densities decreased after 6 days in corals kept in total darkness, although chlorophyll concentrations remained higher. Corals that were fed and subjected to 90% PAR(0) showed lower degrading zooxanthellae frequencies, higher photosynthetic and respiration rates, and higher chlorophyll concentrations than corals in the same light regime under starvation. Complete acclimation to dim (20% PAR(0)) and low (3% PAR(0)) light was only apparent for corals fed with zooplankton. Changes in zooxanthellae population densities occurred through differential rates of zooxanthellae division and degradation.