ESTIMATING CONTRIBUTION OFZOOXANTHELLAE to ANIMAL RESPIRATION (CZAR) and to ANIMAL GROWTH (CZAG) of GIANT CLAM Tridacna Maxima

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ESTIMATING CONTRIBUTION OFZOOXANTHELLAE to ANIMAL RESPIRATION (CZAR) and to ANIMAL GROWTH (CZAG) of GIANT CLAM Tridacna Maxima Journal of Coastal Development ISSN : 1410 - 5217 Volume 9, Number 3, June 2006 : 155-162 Accredited : 23a/Dikti/Kep/2004 ESTIMATING CONTRIBUTION OFZOOXANTHELLAE TO ANIMAL RESPIRATION (CZAR) AND TO ANIMAL GROWTH (CZAG) OF GIANT CLAM Tridacna maxima Ambariyanto *) Marine Science Department, Diponegoro University, Kampus Tembalang, Semarang, 50275, Indonesia. Received : April, 20, 2006 ; Accepted : May 25, 2006 ABSTRACT Zooxanthellae are symbiotic dinoflagellate algae which live in association with marine invertebrates including giant clams. These algae are capable of translocating part of their photosynthetic products to the host. This translocation is one of the nutrition sources of the host. The present study aims to calculate the contribution of zooxanthellae on the energy requirements of adult giant clam (Tridacna maxima) during their respiration and growth processes. The result showed that zooxanthellae are capable of contributing 260.67% and 452.54% energy required by giant clams for respiration and growth during summer and 171.51% and 273.51 % during winter, respectively. It suggests that all the energy required for these two processes can be supplied by zooxanthellae. Key words : giant clams, zooxanthellae, energy, CZAR, CZAG *Corespondence : Phone / fax : 024. 7474 698 ; Email : [email protected], INTRODUCTION Giant clams are marine bivalves which live have been degrading rapidly including in in coral reef ecosystems in the Indo-Pacific Indonesia. region. There are two genera and eight Giant clams are also known as species of giant clams around the world, and marine invertebrate which live in seven of them can be found in Indonesian association with single cell dinoflagelate waters, except Tridacna tevoroa which only algae commonly known as zooxanthellae. Lives in South Pacific countries These algae which can be found in the (Romimohtarto et al., 1987; Knop, 1996). mantle of the clams has an important role as Traditionally, these clams were used by a source of energy for the host. These algae coastal people as a source of meat, building are capable of translocating part of their materials, and other purposes such as house photosynthetic products to the host which hold equipments. Recently, these clams have then be used for biological processes become important export commodities by (Lucas, 1994). several countries mainly for aquarium So far there is no study available animals (Lucas, 1994). Consequently, the especially, on the estimation of the role of condition of giant clams natural populations zooxanthellae on energy requirement by the clams. This study aims at estimating the Estimating Contribution of Zooxanthellae to Animal Respiration (CZAR) and to Animal Growth (CZAG) of Giant Clam, Tridacna maxima 155 Journal of Coastal Development ISSN : 1410 - 5217 Volume 9, Number 3, June 2006 : 155-162 Accredited : 23a/Dikti/Kep/2004 contribution of zooxanthellae to animal energy requirements for respiration (CZAR) Carbon content of zooxanthellae was and for growth (CZAG). calculated using an equation that relates cell volume and elemental carbon (Strathmann, MATERIALS AND METHODS 1967). Cell volume of zooxanthellae was calculated base on the mean size of 100 zooxanthellae isolated from the giant clams. Giant clams, Tridacna maxima were The equation to calculate carbon content is collected from intertidal areas on the reef as follows: crest at One Three Island lagoon, Great log carbon = -0.314+0.712 log cell Barrier Reef, Australia using permit at the volume (Strathmann, 1967) depth of approximately 150 cm. These clams were grown in microatolls during winter and summer months. Isolation of zooxanthellae Carbon required for zooxanthellae growth. was done by using the method used by Carbon used by zooxanthellae for growth Ambariyanto and Hoegh-Guldberg (1997). can be calculated by multiplying In order to calculate CZAR and zooxanthellae specific growth rate and CZAG several parameters are required. carbon content, as the following: These are the specific growth rates of C = µ x Carbon per cell zooxanthellae, the carbon used for Zg zooxanthellae growth and respiration, and Carbon translocated by zooxanthellae the carbon used for animal growth and respiration (Muscatine et al., 1981; Carbon translocated can be calculated by Ambariyanto, 2002). the following formula, T = P - R - G Z Z Z where : Specific growth rate of zooxanthellae. PZ = carbon gained from Zooxanthellae specific growth rate zooxanthellaephotosynthesis was calculated by using an equation first RZ = carbon used for zooxanthellae used by Wilkerson et al. (1983). This respiration formula is based on the mitotic index (f = the GZ = carbon used for zooxanthellae percent of dividing cells in a population) and growth the duration of paired cell stage (td). Mitotic index was calculated based Estimating CZAR and CZAG on the number of dividing zooxanthellae in 1000 isolated zooxanthellae times 100 CZAR can be calculated by the following percent. While the value of td used in this formula : CZAR = T / RA, where calculation is 0.46 d based on several studies T = carbon translocated and (Wilkerson et al., 1983; Muscatine et al., RA = carbon required by the animal 1984; Hoegh-Guldberg et al., 1986). It is for respiration also noted that variation in td has only small While CZAG can be calculated by the impacts on CZAR (Muscatine et al., 1981). following formula : µ = 1/td ln(1+fmax) CZAG = (T - RA.)/Carbon growth where µ = zooxanthellae specific growth rate t = 0.46 (Wilkerson et al., 1983; Muscatine d et al., 1984). fmax= mitotic index. Carbon content of the zooxanthellae. RESULTS AND DISCUSSION Estimating Contribution of Zooxanthellae to Animal Respiration (CZAR) and to Animal Growth (CZAG) of Giant Clam, Tridacna maxima 156 Journal of Coastal Development ISSN : 1410 - 5217 Volume 9, Number 3, June 2006 : 155-162 Accredited : 23a/Dikti/Kep/2004 Specific growth rate of zooxanthellae. Pteraeolidia ianthina; Hoegh-Guldberg et al., 1986), and 88.85 and 86.5 pg cell-1 The result of this study showed that (zooxanthellae from light and shade adapted maximum value of mitotic index is 0.025, coral Stylophora pistillata respectively; thus the specific growth is : Muscatine et al., 1984). Using the average -1 µ = 1/0.46 ln (1+0.025) = 0.054 day .... (1) value (77.7 pg cell-1), the protein content in zooxanthellae is Carbon content of the zooxanthellae. = 3.8 108 cell x 77.7 pg cell-1 = 29.53 mg The results of the present study showed that the cross-sectional area of zooxanthellae in The total protein content of zooxanthellae the clams was 47.9 µm2. The cell radius (r) is and the animal host (giant clam protein hence: content) was calculated by multiplying the r = √47.9/π = 3.9 µm, and the 3 protein content per gram mantle (5.329 mg Cell volume = 4/3 x π x r = -1 3 g ) by the total wet tissue weight (47.311 g; 248.47 µm Ambariyanto, 1996). It is assumed that Carbon content of zooxanthellae can be protein content within clam mantle and calculated as follows other internal parts is similar. Hence, the log carbon = -0.314+0.712 log cell volume -1 total protein content in zooxanthellae and Carbon per cell = 24.628 pg C cell ........(2) animal is = 5.329 mg g-1 x 47.311 g = 252.12 mg Carbon required for zooxanthellae growth. -1 The percent of the total respiration due to CZg = µ x Carbon. cell = 0.0054 day-1 x 24.628 pgC zooxanthellae was -1 =29.53 mg/252.12 = 0.117, or = 11.7 % cell -1 -1 = 1.3299 pgC cell day The mean value of total respiration by intact Carbon required for growth of the entire clam (RC) in this model was 8 -1 -1 population of zooxanthellae (=3.8 10 cell RC= 0.172 mmol O2 clam h clam-1) is (Ambariyanto, 1996), which is 8 -1 - 1 CZg = 3.8 10 cell x 1.3299 pgC = 0.172 x 32 =5.504 mg O2 clam h = -1 -1 -1 -1 cell day 5504 µg O2 clam h -1 -1 - = 0.502 mg C day ...... (3) = 5.504 x 24 = 132.09 mg O2 clam day 1 Respiration of giant clam and This value can be converted into zooxanthellae. carbon using RQA (respiratory quotient for the animal = 0.8; Muscatine et al., 1981) The respiration due to the and the ratio of the masses of C and O2 (ie. zooxanthellae within a clam is a fraction of 12/32 or 0.375), as the following the total respiration. In previous studies the ratio of zooxanthellae and animal biomass -1 -1 RC = 5504 µg O2 clam h x 0.375 x RQ has been used to calculate the portion of the = 5504 µg O clam-1 h-1 x 0.375 x 0.8 total respiration due to zooxanthellae 2 = 1651.2 µg C clam-1 h-1 (Muscatine et al., 1984; Hoegh-Guldberg et -1 -1 al., 1986). = 39.63 mg C clam day .............(4) Protein content per zooxanthellae is The carbon required for 77.7 pg cell-1. This value is based on an zooxanthellae respiration (RZ) was average of values reported from the literature calculated by multiplying (4) by the for similar size of cells. These were 58.1 pg proportion on the total respiration due to the cell-1 (zooxanthellae from nudibranch zooxanthellae (that is 0.117, see above) : Estimating Contribution of Zooxanthellae to Animal Respiration (CZAR) and to Animal Growth (CZAG) of Giant Clam, Tridacna maxima 157 Journal of Coastal Development ISSN : 1410 - 5217 Volume 9, Number 3, June 2006 : 155-162 Accredited : 23a/Dikti/Kep/2004 = 96.9 % -1 -1 -1 RZ = 39.63 mg C clam day x 0.117 = b). Twinter = 60.01 mg C day - 0.502 mg 4.64 mg C day-1 ......................... (5) C day-1 - 2.32 mg C day-1 =57.19 mg C day-1...........................(10) Therefore, carbon required for animal %Twinter= (57.19/60.01) x 100 % = 95.3 % respiration (RA): -1 RA = 39.63 - 4.64 = 34.99 mg C day .
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