Viruses in aquatic ecosystems: important advancements of the last 20 years and prospects for the future in the field of microbial oceanography and limnology

Over the last two decades, viruses in aquatic systems have been observed to modify, influence and control aquatic systems. Since the determination decades ago that viruses were abundant in aquatic ecosystems, researchers have demonstrated that viruses are pervasive and dynamic across the expanse and depth of all aquatic systems as well as at the water-sediment interface. There have been a wide range of methodological advancements during this time. To date, aquatic viruses have been suggested to play vital roles in global and small-scale biogeochemical cycling, community structure, algal bloom termination, gene transfer, and evolution of aquatic organisms. Even in harsh and difficult to study environments, aquatic and benthic viruses have been demonstrated to be major players in carbon cycling and recycling of nutrients from organic material. Taxonomic and metagenomic research has shown us that there are major globally-distributed groups, but that their genomes are filled with sequence information that has no similarity to sequences in existing bioinformatic databases. And while the field of viral ecology has expanded exponentially since the late 1980s, there is much that we do not yet understand about virusmediated processes in aquatic systems. Important near-term steps include the combination of advanced metagenomic techniques with studies of function and population control, standardization of methodological approaches to facilitate global data acquisition without concern over methods-based artefacts, understanding of viral life strategies and their triggers, and the role of viruses in the transformation of organic matter. The purpose of this manuscript is to bring the reader a review of the recent advances in aquatic viral ecology in light of new areas of research, applications of viral ecology to real-world problems, and refinement of models of viral interactions on a range of scales.