Effect of nutrient enrichment in the field on the biomass, growth and calcification of the giant clam Tridacna maxima

Nutrients were added separately and combined to an initial concentration of 10 μM (ammonium) and/or 2 μM (phosphate) in a series of experiments carried out with the giant clam Tridacna maxima at 12 microatolls in One Tree Island lagoon, Great Barrier Reef, Australia (ENCORE Project). These nutrient concentrations remained for 2 to 3 h before returning to natural levels. The additions were made every low tide (twice per day) over 13 and 12 mo periods for the first and second phase of the experiment, respectively. The nutrients did not change the wet tissue weight of the clams, host C:N ratio, protein content of the mantle, calcification rates or growth rates. However, ammonium (N) enrichment alone significantly increased the total population density of the algal symbiont (Symbiodinium sp.: C = 3.6 · 10⁸ cell clam⁻¹, N = 6.6 · 10⁸ cell clam⁻¹, P = 5.7 · 10⁸ cell clam⁻¹, N + P = 5.7 · 10⁸ cell clam⁻¹; and C = 4.1 · 10⁸ cell clam⁻¹, N = 5.1 · 10⁸ cell clam⁻¹, P = 4.7 · 10⁸ cell clam⁻¹, N + P = 4.5 · 10⁸ cell clam⁻¹, at the end of the first and second phases of the experiment, respectively), although no differences in the mitotic index of these populations were detected. The total chlorophyll a (chl a) content per clam but not chlorophyll a per cell also increased with ammonium addition (C = 7.0 mg chl a clam⁻¹, N = 13.1 mg chl a clam⁻¹, P = 12.9 mg chl a clam⁻¹, N + P = 11.8 mg chl a clam⁻¹; and C = 8.8 mg chl a clam⁻¹, N = 12.8 mg chl a clam⁻¹; P = 11.2 mg chl a clam⁻¹, N + P = 11.3 mg chl a clam⁻¹, at the end of the first and second phases of the experiment, respectively). The response of clams to nutrient enrichment was quantitatively small, but indicated that small changes in inorganic nutrient levels affect the clam–zooxanthellae association. Received: 2 June 1997 / Accepted: 9 June 1997     

Influence of field-based nutrient enrichment on the photobiology of the giant clam Tridacna maxima

Nutrients were added to 12 microatolls in One Tree Island lagoon every low tide for 13 mo to an initial concentration of 10 μM (ammonium, N) and 2 M (phosphate, P). These concentrations remained above background for 2 to 3 h after addition. The addition of ammonium (N and N+P but not P alone) significantly increased P _g (gross photosynthesis) P _n (net photosynthesis) and R (respiration) per unit wet-tissue weight and α (photosynthetic efficiency) in Tridacna maxima after 3 mo nutrient enrichment. These responses to small and transient changes in ammonium concentrations suggest that symbiotic clams are not nutrient-replete, and that even subtle changes in nutrients can have a measurable effect on photosynthesis. The same clams did not show significant differences in photosynthetic parameters 6 mo after the beginning of nutrient enrichment, suggesting that their previous responses had either been seasonal or that symbiotic clams such as T. maxima are able to adjust their photophysiology following external changes in nutrient concentrations. Received: 26 August 1997 / Accepted: 11 December 1998     

Nutrient enrichment and the ultrastructure of zooxanthellae from the giant clam Tridacna maxima

The separate and combined effects of ammonium (10M) and phosphate (2M) on the ultrastructure of zooxanthellae (Symbiodinium sp.) from giant clams, Tridacna maxima, were examined in the field. Nitrogen addition significantly changed the ultrastructure of the zooxanthellae inhabiting the clams. After 9 mo exposure, the cross-sectional area of zooxanthellae from N-treated clams was significantly lower than that from other treatments [N=39.3 m²; C=47.9 m²; P=43.2m²; N+P=44.5 m²; (P=0.001)]. There was also a significant decrease in the size of starch bodies, especially around the pyrenoid of the zooxanthellae from N and N+P treatments [N=1.2 m²; C=2.0 m²; P=1.8 m²; N+P=1.2 m²; (P=2.08E-11)]. This presumably occurs as a result of the mobilization of organic carbon stores in response to stimulated amino acid synthesis under enriched nutrient conditions. These data strongly suggest that the symbiotic zooxanthellae of clams are limited to some extent by the availability of inorganic nitrogen, and that relatively minor changes to the nutrient loading of the water column can have substantial effects on the biochemistry of symbioses such as that which exists between clams and zooxanthellae.