New Insights Into the Virus-to-Prokaryote Ratio (VPR) in Marine Sediments

The virus-to-prokaryote ratio (VPR), which reflects the numerical dominance of viruses over their hosts, has been proposed as a proxy for assessing the relationship between viruses and prokaryotes. Previous studies showed that VPR values fluctuate over six orders of magnitude within and across various benthic ecosystems, with an average value of approximately 10. We hypothesize that this high VPR value is largely due to the inaccurate enumeration of viruses and prokaryotes (e.g., centrifugation treatments may lead to a three–fourfold overestimation of VPR). In this study, we evaluated the impact of processing methods on the determination of VPR values. Using an optimized procedure, we investigated the marine benthic VPR at 31 sites, from intertidal zones through continental shelves to abyssal plains, and assessed its monthly variation in two contrasting intertidal habitats (muddy-sand and sandy). By compiling 135 VPR data points of surface sediments from 37 publications, we reveal the effect of centrifugation on published VPR values and describe the spatial distribution of VPR values on a larger scale based on reliable data. The results showed that the commonly used centrifugation method may result in an overestimation of VPR values that are approximately one order of magnitude higher than those obtained using the dilution method, while other processing steps had a limited impact on the VPR. Our analysis indicates that the benthic VPR value is low and less varied across temporal and spatial scales, fluctuating mostly within 10, and the average VPR is approximately 2 in both marine and freshwater habitats. An insignificant seasonal pattern in the VPR was observed in the intertidal zone, with lower VPR values occurring at high temperatures. The VPR spatial distribution was primarily associated with sediment phaeophytin a, suggesting that the trophic conditions of the upper water column and the sedimentation of organic matter to the bottom are the key factors affecting VPR values. The mean VPR in benthic habitats is approximately one order of magnitude lower and much less varied than that observed in pelagic habitats, indicating that the virus–host relationship and the ecological function of viruses in the two ecosystems may be very different.