Viruses in multi-scale ocean models: challenges and opportunities

While viruses affect the flow of elements and energy at a planet-wide scale through lysis, gene transfer, and metabolic reprogramming, they are yet to be included in planetary-scale models of ecosystem function and nutrient cycling. Here, we review recent advances incorporating viruses into ocean models and ask: what barriers remain? To address these challenges, we argue for a new generation of ocean models that are fully representative of the multifaceted influences of viruses across scales of organization. We describe ways to achieve this by integration of existing models built across scales, from molecules to ecosystems and the Earth System. To accelerate these advances, we emphasize the need for systematic, intercalibrated datasets for diverse experimental virus-host systems, wider application of new technologies to monitor in situ viral infections, and new software to integrate models across scales. Resolution of viruses within multi-scale models will open the door to assessing current biological uncertainties related to the impact of viral infection on nutrient retention in the surface ocean, carbon sequestration to depth, and the sensitivity of these processes to climate change.