A light-dependent ammonia-assimilating mechanism in the ctenidia of a giant clam

Giant clams harbor symbiotic zooxanthellae and manifest light-enhanced calcification. In light, they also increase the absorption and assimilation of exogenous ammonia, but the roles of the host and symbionts are unclear and the reason for light dependency remains enigmatic. Here, we report the complete coding cDNA sequence of a glutamine synthetase (GS), from the ctenidia (gill) of Tridacna squamosa. The deduced GS amino acid sequence was closely related to other animal GSs, indicating a host origin. The GS/GS transcript level and protein abundance increased significantly in the ctenidia after 12 h of light exposure. These results denote the ctenidia as the site of light-enhanced ammonia absorption with the absorbed ammonia being assimilated to glutamine by the host GS. Immunofluorescence microscopy confirmed that GS was expressed predominantly in the cytoplasm of the epithelial cells separating the tertiary water channels from the hemolymph inside the ctenidia. Hence, unlike symbiotic cnidarians, the host clam probably supplies exogenous nitrogen as glutamine from the ctenidia, through the hemolymph and tubular fluid, to the nitrogen-deficient zooxanthellae which reside extracellularly in the extensible outer mantle. This is the first report on light-dependent gene and protein expression of a host enzyme involved in nitrogen metabolism in an alga–invertebrate association