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Experimental Manipulations of a Solitary Coral (<I BULLETIN OF MARINE SCIENCE, 56(1): 319-329, 1995 CORAL REEF PAPER EXPERIMENTAL MANIPULATIONS OF A SOLITARY CORAL (FUNGI A, SCLERACTINIA) WITH EMPHASIS ON THE EFFECTS OF LIGHT Helen T. Yap, A. Rex F. Montebon, f.-A. von Oertzen and Romeo M. Dizon ABSTRACT The metabolic rate per unit area of the solitary scleractinian coral Fungia (Danafungia) horrida did not vary significantly within a size range of 5-14 cm (diameter), Likewise. maintenance time in the laboratory up to a period of 1 week did not appear to influence metabolic performance. Gross photosynthetic rates, when measured under similar light inten- sities, did not differ significantly between corals in the field and in the laboratory. Significant variations in photosynthetic response to light occurred. Combined data for this and other species of the genus Fungia from the same reef flat were used to generate light response curves which showed I, to be about 800 microEinsteins m-2 S-I, as determined from the hyperbolic tangent formulation. Theoretical maximum gross primary production was about 2 I 250 fJ.g O2 cm- h- • Primary production and respiration rates in scleractinian corals have not been measured previously in the Philippines, although a large body of literature on the subject exists for many geographic locations (reviewed in McCloskey et aI., 1978; Gladfelter, 1985; Sebens, 1987). Laboratory experiments on corals have dealt mostly with responses of various species to light. Examples are the investigations of Zvalinskii et aI. (1980), Chalker et aI. (1983), Kinzie and Hunter (1987), Den- nison and Barnes (1988), Edmunds and Davies (1988, 1989), and Newton and Atkinson (1991). Knowledge of light responses of corals is crucial in the interpretation of data which are eventually used in energy budgets. The coral-zooxanthellae complex responds instantly to changes in light intensity. There are particular ranges of light intensity which are optimal for primary production for each species (Sebens, 1987). Below such ranges, gross photosynthetic rate decreases and is eventually exceeded by respiration. Above optimal light intensities, the photosynthetic ma- chinery becomes saturated. Light saturation curves are especially useful for the investigation of such responses (Zvalinskii et aI., 1980; Chalker, 1980, 1981; Wyman et aI., 1987). One such curve is constructed here for the genus Fungia. There appears to be a dearth of studies to date on the effects of maintenance time, size, and physical transfer to the laboratory on coral metabolic rates. Because of logistic constraints that often arise, a number of metabolic experi- ments are done in the laboratory, and their results extrapolated to in situ condi- tions. Or at least, such results are presumed to apply to in situ conditions. It is with these constraints in mind that the effects of transfer to the laboratory and of maintenance time there are examined in this study. Finally, the question of size as it influences metabolic activity is still an open question for many organisms, including corals. Growth, for example, may be faster in smaller colonies (Bud- demeier and Kinzie, 1976; Yap et aI., 1992), although this appears not to be a general rule (Maragos, 1978; Kinzie and Sarmiento, 1986). In a review of coelen- terate biology, Sebens (1987) reports that weight-specific oxygen consumption decreases with increasing individual size, although his data are compiled mostly from studies on sea anemones. In the case of scleractinian corals, size is probably 319 320 BULLETIN OF MARINE SCIENCE. VOL. 56, NO. I. 1995 a factor that influences metabolic rates. This, again, has implications for conclu- sions that are drawn, i.e., the resulting energy budgets may be valid for only a particular size range of the organisms in question. In this paper, the results of experiments on various aspects of metabolism are described. These experiments address the following questions: ]) Are there sig- nificant changes in metabolic rate with respect to length of time maintained in the laboratory (from collection in the field) up to a period of 1 week? 2) Are there significant differences in metabolic rates with respect to size (within a certain range) of the corallum? 3) What is the photosynthetic response of the coral- zooxanthellae complex to light intensity within the range of natural daylight en- countered? 4) Are there significant differences between metabolic rates measured in situ and in the :laboratory? Metabolic rates here are expressed per unit surface area of the corallum which is a relatively useful standard (McCloskey and Muscatine, 1984). "Production" as used in the text refers to gross primary production or gross photosynthesis, except where light saturation curves were generated, in which case net primary production or net photosynthesis was used instead (Chalker et aI., 1983). MATERIALS AND METHODS Study Sile.-The study site is a 0.5 lan-wide reef flat on the western side of Santiago Island in Bolinao, Pangasinan in the northwestern Philippines at approximately 16°24'41"N lat and 119°54'25"E long. Water depth averages about 1 m. A sea grass zone stretches from the shore about halfway across the flat, where it ends abruptly and gives way to a sand-rubble substrate. This second zone is dominated by large microatolls of Porites. Specimens of the solitary coral Fungia (Danafungia) horrida were collected from this area. The reason fungiids were ~;elected for study is that they consist of a single polyp with relatively few endoliths, rendering the interpretation of physiological results easier. In addition to the coral animal itself and the symbiotic zooxanthellae, coral colonies, in contrast, are inhabited by a variety of com- mensal or parasitic plants and animals which also contribute to the observed oxygen changes. Mea- surements from a single polyp such as F. (Danafungia) horrida or other similar species of Fungia are therefore more straightforward, and can readily be attributed mainly to the coraI-zooxanthellae com- plex. Metabolic Experiments.-The metabolic experiments performed in this study made use of the oxygen technique (Zeitzschel, 1981). Primary production and respiration were measured by the degree of oxygen evolution or consumption, respectively. Metabolic rates are expressed per unit surface area of the coral. More specifically, projected surface area was used (Yap et aI., 1992). This was obtained by measuring the perceived longest dimension of the coral through the center (length, L) with a caliper calibrated in millimeters, and then taking the perceived longest dimension normal to L (width, W). Projected area, A, was computed by using the formula: Field measurements of metabolic rates made use of a Plexiglas cylinder (volume = approx. 9 liters) bolted onto an acrylic platform on which the specimen was placed, forming a watertight enclosure. The top end of the cylinder was fitted with a dissolved oxygen probe (Nester Instruments model 8500X) and a battery-operated bilge pump for stirring the enclosed water (turn-over rate of the pump was about 400 gallons per hour). Prior to a series of measurements during a day, the probe was air calibrated according to the manufacturer's instructions. Respiration was measured first by covering the set-up with a black canvass material to exclude light and, hence, prevent photosynthesis. Production was then measured with the cover removed. Starting with the dark incubation reduced oxygen levels, thus avoiding supersaturation during photosynthetic measurements. For both production and respiration, changes in oxygen were recorded every minute until enough data points were obtained to produce a significant regression. Typically, measurement runs lasted lO- IS min, and took place: between 0900-1500. Temperature inside the chamber was monitored during the entire incubation by the probe's temper- ature sensor. This was compared with temperature outside the chamber which was measured with a YAP ET AL.: EXPERIMENTAL MANIPULATIONS OF A SOLITARY CORAL 321 mereury thermometer against which the probe sensor had been ealibrated in the laboratory. In general, internal chamber temperatures were higher than external temperatures by less than 1°C. Light values within the photosynthetically active range of radiation were registered simultaneously by aLI-COR 1935A spherical quantum sensor attached to a data logger. The sensor was mounted onto a tripod support and was situated about I m below the water surface. A spherical type was selected for purposes of measuring ambient light from all directions as a coral would typically receive it. Several light readings were averaged per minute for the entire duration of a production run, yielding a mean light intensity corresponding to a particular photosynthetic rate. A similar procedure was followed in the laboratory. The same enclosure and D.O. meter as in the field experiments were used for "Large" specimens (see explanation of size below). For "Small" and "Medium" corals, the incubation chamber was an acrylic cylinder (Strathkelvin Instruments RC400) with a volume of approximately 750 ml. Stirring was accomplished by means of a magnetic stir bar on the bottom. Dissolved oxygen was measured using a Strathkelvin D.O. meter (Model 781b) fitted with a Clarke-type microelectrode and connected to a stripchart recorder. Prior to a series of mea- surements, the electrode was calibrated as follows: the zero setting was determined after immersing the probe in a sodium borate/sodium sulfite solution to eliminate oxygen. The 100% oxygen saturation level was set by vigorously aerating seawater for at least 20 min, and then immersing the probe in this after thorough rinsing with distilled water. The temperature and salinity of the medium in both types of enclosures were kept at the average levels characterizing field conditions during most of the year. Salinity was 34%0, and temperature was fixed at about 28°C. Light intensity in the laboratory was regulated by a fluorescent light bank which could attain intensities of up to 400 microEinsteins m-2 s-'. A pair of halogen lamps (1,000 W each) was used to achieve intensities up to 750 f1E m-2 s-' which more elosely approximated field conditions during most daylight hours.
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