Compared to our understanding of the taxonomic composition of the symbiotic microbes in marine sponges, the functional diversity of these symbionts is largely unknown. Furthermore, the application of genomic, transcriptomic, and proteomic techniques to functional questions on sponge host-symbiont interactions is in its infancy. In this study, we generated a transcriptome for the host and a metatranscriptome of its microbial symbionts for the giant barrel sponge, Xestospongia muta, from the Caribbean. In combination with a gene-specific approach, our goals were to (1) characterize genetic evidence for nitrogen cycling in X. muta, an important limiting nutrient on coral reefs (2) identify which prokaryotic symbiont lineages are metabolically active and, (3) characterize the metabolic potential of the prokaryotic community. Xestospongia muta expresses genes from multiple nitrogen transformation pathways that when combined with the abundance of this sponge, and previous data on dissolved inorganic nitrogen fluxes, shows that this sponge is an important contributor to nitrogen cycling biogeochemistry on coral reefs. Additionally, we observed significant differences in gene expression of the archaeal amoA gene, which is involved in ammonia oxidation, between coral reef locations consistent with differences in the fluxes of dissolved inorganic nitrogen previously reported. In regards to symbiont metabolic potential, the genes in the biosynthetic pathways of several amino acids were present in the prokaryotic metatranscriptome dataset but in the host-derived transcripts only the catabolic reactions for these amino acids were present. A similar pattern was observed for the B vitamins (riboflavin, biotin, thiamin, cobalamin). These results expand our understanding of biogeochemical cycling in sponges, and the metabolic interchange highlighted here advances the field of symbiont physiology by elucidating specific metabolic pathways where there is high potential for host-prokaryote interactions.

Excess nitrogen (N) loading and resulting eutrophication plague coastal ecosystems globally. Much work is being done to remove N before it enters coastal receiving waters, yet these efforts are not enough. Novel techniques to remove N from within the coastal ecosystem are now being explored. One of these techniques involves using oysters and their habitats to remove N via denitrification. There is substantial interest in incorporating oyster-mediated enhancement of benthic denitrification into N management plans and trading schemes. Measuring denitrification, however, is expensive and time consuming. For large-scale adoption of oystermediated denitrification into nutrient management plans, we need an accurate model that can be applied across ecosystems. Despite significant effort to measure and report rates of denitrification in oyster habitats, we are unable to create such a model, due to methodological differences between studies, incomplete data reporting, and inconsistent measurements of environmental variables that may be used to predict denitrification. To make a model that can predict denitrification in oyster habitats a reality, a common sampling and reporting scheme is needed across studies. Here, we provide relevant background on how oysters may stimulate denitrification, and the importance of oyster-mediated denitrification in remediating excess N loading to coastal systems. We then summarize methods commonly used to measure denitrification in oyster habitats, discuss the importance of various environmental variables that may be useful for predicting denitrification, and present a set of guidelines for measuring denitrification in oyster habitats, allowing development of models to support incorporation of oyster-mediated denitrification into future policy decisions.The past 20 yr have seen rapid development of the oyster aquaculture industry (Fig. 1) and substantial oyster reef restoration (Bersoza Hern andez et al. 2018;Duarte et al. 2020). In this same time period, there has been a large research effort to quantify whether-and by how much-oysters enhance benthic denitrification (the conversion of biologically available dissolved nitrogen [N] to inert di-nitrogen [N 2 ] gas). Early suggestions of the potential for enhanced denitrification in oyster habitats piqued the interest of ecologists, coastal managers, and oyster farmers who imagined a new N mitigation tool that could be included in nutrient management plans (Newell

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