Succession of free-living and particle associated prokaryotes in the northern Adriatic Sea

Marine snow is composed of a complex pool of organic and inorganic matter. Microbial communities thrive in it, using a large number of compounds as a source of energy and nutrients. Microbial abundance and activity have been reported to be orders of magnitude higher in marine snow than in the ambient water. However, it is still unclear whether the microbial community colonizing marine snow comprises mainly generalists or specialists. In this study, we monitored the prokaryotic community inhabiting marine snow (MS) and ambient water (AW) in the northern Adriatic Sea. MS was collected with syringes during SCUBA diving and AW was sampled with Niskin bottles. In the AW, inorganic N:P ratios averaged 30.5 ± 24.8 indicating P-limitation while in MS inorganic N:P ratios were 11 ± 11.9. Prokaryotic abundance in MS was similar to AW, with an enrichment factor (EF, i.e. ratio between MS/AW) of 1.4 ± 1.8. However, the microbial activity was generally higher in MS, extracellular enzymatic activity exhibited EFs ranging from 2.6 ± 1.4 to 8.3 ± 7.7, and the EF of heterotrophic leucine incorporation, a proxy for heterotrophic biomass production, was 7.8 ± 4.9. The relative abundance of different phylogenetic groups in MS changed over the course of the summer, e.g. Sphingobacteriales (from 10.6% to 0.9%), Verrucomicrobia (0.3% to 3.5%) and Actinobacteria (1.2% to 5.5%), indicating successional changes linked to changes in the environmental conditions in MS. The bacterial community inhabiting MS was characterized by a larger relative abundance of Cyanobacteria as compared to AW (27.5% in MS vs. 13.5% in AW) and Planctomycetes (4.8% in MS vs. 1.1% in AW) and a lower abundance of Alphaproteobacteria (20% in MS vs. 38.5% in AW). Taken together, our results indicate the presence of a core bacterial community in MS and AW. The highly active MS community is characterized by the presence of specialized groups that can reach high abundances, and a dynamic generalist community thriving in this nutrient-enriched microenvironment.