The"biological pump",which is based on the gravity-induced vertical sinking of organic carbon,is one of the primary mecha-nisms for carbon storage in the ocean[1].Via this mechanism,atmospheric CO2 fixed by phytoplankton as particulate organic car-bon(POC)is transferred from the surface to the deep sea for long-term sequestration.The amount of carbon flux entering the ocean interior is estimated to be approximately 5-10 Gt per year in the form of POC,which represents a major carbon source for deep-sea ecosystems and plays a crucial role in global marine biogeo-chemical cycles and climate change[1].

It is commonly recognized that viruses control the composition, metabolism, and evolutionary trajectories of prokaryotic communities, with resulting vital feedback on ecosystem functioning and nutrient cycling in a wide range of ecosystems. Although the deep biosphere has been estimated to be the largest reservoir for viruses and their prokaryotic hosts, the biology and ecology of viruses therein remain poorly understood.The deep virosphere is an enigmatic field of study in which many critical questions are still to be answered.Is the deep virosphere simply a repository for deeply preserved, non-functioning virus particles? Or are deep viruses infectious agents that can readily infect suitable hosts and subsequently shape microbial populations and nutrient cycling? Can the cellular content released by viral lysis, and even the organic structures of virions themselves, serve as the source of bioavailable nutrients for microbial activity in the deep biosphere as in other ecosystems? In this review, we synthesize our current knowledge of viruses in the deep biosphere and seek to identify topics with the potential for substantial discoveries in the future.

Marine organisms perform a sea of diel rhythmicity. Planktonic diel dynamics have been shown to be driven by light, energy resources, circadian rhythms, and the coordinated coupling of photoautotrophs and heterotrophic bacterioplankton. Here, we explore the diel fluctuation of viral production and decay and their impact on the total and active bacterial community in the coastal and open seawaters of the South China Sea. The results showed that the night-production diel pattern of lytic viral production was concurrent with the lower viral decay at night, contributing to the accumulation of the viral population size during the night for surface waters. The diel variations in bacterial activity, community composition, and diversity were found highly affected by viral dynamics. This was revealed by the finding that bacterial community diversity was positively correlated to lytic viral production in the euphotic zone of the open ocean but was negatively related to lysogenic viral production in the coastal ocean. Such distinct but contrasting correlations suggest that viral life strategies can not only contribute to diversifying bacterial community but also potentially piggyback their host to dominate bacterial community, suggesting the tightly synchronized depth-dependent and habitat-specific diel patterns of virus–host interactions. It further implies that viruses serve as an ecologically important driver of bacterial diel dynamics across the ocean, highlighting the viral roles in bacterial ecological and biogeochemical processes in the ocean.

Groundwater ecosystems are typically poor in organic carbon and productivity sustaining a low standing stock of microbial biomass. In consequence, microbial food webs in oligotrophic groundwater are hypothesized to be bottom-up controlled. To date, quantitative information on groundwater microbial communities, food web interactions, and carbon flow is relatively lacking in comparison to that of surface waters. Studying a shallow, porous alpine aquifer we collected data on the numbers of prokaryotes, virus-like particles and heterotrophic nanoflagellates (HNFs), the concentration of dissolved (DOC) and assimilable organic carbon (AOC), bacterial carbon production (BCP), and physical-chemical conditions for a 1 year hydrological cycle. The potential effects of protozoan grazing and viral lysis onto the prokaryotic biomass was tested. Flow of organic carbon through the microbial food web was estimated based on data from the literature. The abundance of prokaryotes in groundwater was low with 6.1 ± 6.9 × 104 cells mL–1, seasonally influenced by the hydrological dynamics, with higher densities coinciding with a lower groundwater table. Overall, the variability in cell numbers was moderate, and so it was for HNFs (179 ± 103 HNFs mL–1) and virus-like particles (9.6 ± 5.7 × 105 VLPs mL–1). The virus to prokaryotes and prokaryote to HNF ratios ranged between 2–230 and 33–2,084, respectively. We found no evidence for a viral control of prokaryotic biomass, and the biomass of HNFs being bottom-up controlled. First estimations point at carbon use efficiencies of 0.2–4.2% with prokaryotic production, and carbon consumed and recycled by HNFs and phages to be of minor importance. This first groundwater microbial food web analysis strongly hints at a bottom-up control on productivity and standing stock in oligotrophic groundwater ecosystems. However, direct measurement of protozoan grazing and phage mediated lysis rates of prokaryotic cells are urgently needed to deepen our mechanistic understanding. The effect of microbial diversity on the population dynamics still needs to be addressed.

Particle sinking is an important process in the ocean, influencing the biogeochemical cycle and driving the long-term preservation of carbon into the deep sea via the biological pump. However, as an important component of marine ecosystems, the role of viruses during sinking is still poorly understood. Therefore, we performed a series of transplantation experiments in the South China Sea to simulate environmental changes during sinking and investigate their effects on viral eco-dynamics and life strategy. Our study demonstrated increased viral production but decreased virus-mediated bacterial mortality after transplantation. A larger burst size and switch from the lysogenic to lytic strategy were shown to contribute to enhanced viral productivity. We provide experimental evidence that surface viral ecological characteristics changed dramatically after transplantation into deep-sea waters, indicating a potential importance of viruses during vertical sinking in the ocean. This effect probably provides positive feedback on the efficiency of the biological pump.

The viral component in aquatic systems clearly needs to be incorporated into future ocean and inland water climate models. Viruses have the potential to influence carbon and nutrient cycling in aquatic ecosystems significantly. Changing climate likely has both direct and indirect influence on virus-mediated processes, among them an impact on food webs, biogeochemical cycles and on the overall metabolic performance of whole ecosystems. Here we synthesise current knowledge on potential climate-related consequences for viral assemblages, virus-host interactions and virus functions, and in turn, viral processes contributing to climate change. There is a need to increase the accuracy of predictions of climate change impacts on virusdriven processes, particularly of those linked to biological production and biogeochemical cycles. Comprehension of the relationships between microbial/viral processes and global phenomena is essential to predict the influence on as well as the response of the biosphere to global change.

Phytoplankton contribute almost half of the world's total primary production. The exudates and viral lysates of phytoplankton are two important forms of dissolved organic matter (DOM) in aquatic environments and fuel heterotrophic prokaryotic metabolism. However, the effect of viral infection on the composition and biological availability of phytoplankton-released DOM is poorly understood. Here, we investigated the optical characteristics and microbial utilization of the exudates and viral lysates of the ecologically important unicellular picophytoplankton Prochlorococcus Our results showed that Prochlorococcus DOM produced by viral lysis (Pro-vDOM) with phages of three different morphotypes (myovirus P-HM2, siphovirus P-HS2, and podovirus P-SSP7) had higher humic-like fluorescence intensities, lower absorption coefficients, and higher spectral slopes than DOM exuded by Prochlorococcus (Pro-exudate). The results indicate that viral infection altered the composition of Prochlorococcus-derived DOM and might contribute to the pool of oceanic humic-like DOM. Incubation with Pro-vDOM resulted in a greater dissolved organic carbon (DOC) degradation rate and lower absorption spectral slope and heterotrophic bacterial growth rate than incubation with Pro-exudate, suggesting that Pro-vDOM was more bioavailable than Pro-exudate. In addition, the stimulated microbial community succession trajectories were significantly different between the Pro-exudate and Pro-vDOM treatments, indicating that viral lysates play an important role in shaping the heterotrophic bacterial community. Our study demonstrated that viral lysis altered the chemical composition and biological availability of DOM derived from Prochlorococcus, which is the numerically dominant phytoplankton in the oligotrophic ocean.IMPORTANCE The unicellular picocyanobacterium Prochlorococcus is the numerically dominant phytoplankton in the oligotrophic ocean, contributing to the vast majority of marine primary production. Prochlorococcus releases a significant fraction of fixed organic matter into the surrounding environment and supports a vital portion of heterotrophic bacterial activity. Viral lysis is an important biomass loss process of Prochlorococcus However, little is known about whether and how viral lysis affects Prochlorococcus-released dissolved organic matter (DOM). Our paper shows that viral infection alters the optical properties (such as the absorption coefficients, spectral slopes, and fluorescence intensities) of released DOM and might contribute to a humic-like DOM pool and carbon sequestration in the ocean. Meanwhile, viral lysis also releases various intracellular labile DOM, including amino acids, protein-like DOM, and lower-molecular-weight DOM, increases the bioavailability of DOM, and shapes the successive trajectory of the heterotrophic bacterial community. Our study highlights the importance of viruses in impacting the DOM quality in the ocean.

The basic bacterial ecology and diversity was investigated in five running water systems of Southern New Caledonia. These running waters were characterized by potential P-limitation and high concentrations of Ni, Fe, Mn, Cr and Co. The low concentrations of dissolved organic carbon, bacterial and viral abundance, bacterial production and growth efficiency support the characterization of the running waters as oligotroph to ultraoligotroph. Despite these similarities, there were strong differences (<50% similarity) in bacterial community composition between some habitats based on 16S rRNA gene and denaturing gradient gel electrophoresis (DGGE) fingerprints. The high coverage of sequenced DGGE bands found for Betaproteobacteria is typical for freshwater systems, however, we found also a strong representation of Gammaproteobacteria. Indeed the three bands found at all stations were related to Limnohabitans (Comamonadaceae) and Alteromonadaceae. Strong differences were also found between the free-living and the attached bacterial fraction with Gammaproteobacteria dominating in two systems. A higher representation of Gammaproteobacteria seems typical for metal-rich freshwater habitats. Consistent with fresh water habitats, majority of phylotypes detected in the sediment was affiliated to proteobacteria. Also, none of the sequences showed a 100% identity with data bases, and 10 of the 22 and 2 of the 23 sequences had similarities higher than 97% in the freshwater and sediment. This could indicate specific adaptations of the community composition either due to the high metal concentrations or due to the geographical isolation of the New Caledonia.

Microbial communities on marine aggregates could be influenced by ambient water conditions; however, empirical data are scarce. In this study, we used fingerprint analysis of PCR-amplified 16S rRNA gene fragment to examine how microbial communities on aggregates change in response to different conditions of ambient water. We conducted two experiments using seawater cultures from surface waters of the lagoon and the anthropogenically influenced bay of Nouméa, New Caledonia: a transplant experiment in which the artificially produced aggregates from one station was added to ultra-filtered seawater culture of another station, and a water-flow experiment in which the artificially produced aggregates placed in the ultra-filtered seawater culture with or without water-flow. In a transplant experiment, bacterial community composition (BCC) on the bay and lagoon water aggregates were significantly different (p < 0.05, ANOSIM) at the beginning of experiment. After 11 days of incubation, BCC on the lagoon water aggregates were significantly different (p < 0.05) from transplanted communities. Transplantation effect was also observed in the bay water treatments. In a water-flow experiment, BCC on the bay and lagoon water aggregates were significantly different (p < 0.05) at the beginning of the experiment. BCC on the lagoon and bay water aggregates with and without water-flow treatments were significantly different (p < 0.05) at the end of incubation, and effect of water-flow on BCC were observed in the bay and lagoon water treatments. Our experimental studies suggest that changes in ambient water conditions potentially influence microbial communities on aggregates in the Bay of Nouméa.

Viral abundance was assessed in different water masses of the NW Atlantic, and the development of viral abundance, lytic viral infection and lysogeny was followed for the first ca. 5000 km (corresponding to ca. 50 yr in the oceanic conveyor belt) of the western branch of the North Atlantic Deep Water (NADW). Viral abundance was significantly higher in the 100 m layer than in the NADW (2400-2700 m depth) and the Denmark Strait Overflow Water (2400-3600 m depth). The virus-to-prokaryote ratio (VPR) increased with depth, ranging from 32-43 for different water masses of the bathypelagic ocean, thus corroborating the enigma of high viral abundance in the dark ocean. The O2-minimum layer (250-600 m) also showed high viral abundance and VPRs. Viral abundance, a viral subgroup and VPRs decreased in a non-linear form with distance from the NADW origin. Viral production (range: 0.2-2.4 × 107 viruses l-1) and the fraction of lytically infected cells (range: 1-22%) decreased with increasing distance from the formation site of the NADW. Conservative estimations of virus-mediated mortality of prokaryotes in the NADW averaged 20 ± 12%. The fraction of the prokaryotic community with lysogens (i.e. harboring a functional viral DNA) in the NADW averaged 21 ± 14%. Hence, we conclude that (1) viral abundance and subgroups differ between water masses, (2) virus-mediated mortality of prokaryotes as well as lysogeny are significant in the dark ocean and (3) the lysogenic life strategy became more important than the lytic life style during the early formation of the NADW.

The barrier reef lagoon of Noumea (New Caledonia) is potentially influenced by Ni and black carbon (BC) rich aerosols (the two latter originating from a Ni power plant). Experiments were performed with lagoon water collected close to the reef crest to assess the effect of anthropogenic influences on bacterial abundance (BA), production (BP), respiration (BR), growth efficiency (BGE) and community composition (BCC) in artificial produced aggregates. In both treatments, BA were not affected or enhanced compared to the unamended control (up to 40% for BA and 180% for VA). BP was not or negatively affected (up to 70%). No clear trend was found for BR in the Ni treatment; in the BC treatment BR was enhanced by 63-69%. BGE was reduced in both treatments. The strongest effects on BCC profiles as determined by 16S rDNA denaturing gradient gel electrophoresis were due to incubation time treatment particularly for BC. A phylotype corresponding to a specific BC band was closely related to Acinetobacter oleivorans DR1. Thus, this experimental study confirms potential anthropogenic influences on the bacterial community on aggregates in the Bay of Noumea.

In 2005/2006, a major volcanic eruption buried faunal communities over a large area of the 9°N East Pacific Rise (EPR) vent field. In late 2006, we initiated colonization studies at several types of post eruption vent communities including those that either survived the eruption, re-established after the eruption, or arisen at new sites. Some of these vents were active whereas others appeared senescent. Although the spatial scale of non-paved (surviving) vent communities was small (several m2 compared to several km2 of total paved area), the remnant individuals at surviving active and senescent vent sites may be important for recolonization. A total of 46 meio- and macrofauna species were encountered at non-paved areas with 33 of those species detected were also present at new sites in 2006. The animals living at non-paved areas represent refuge populations that could act as source populations for new vent sites directly after disturbance. Remnants may be especially important for the meiofauna, where many taxa have limited or no larval dispersal. Meiofauna may reach new vent sites predominantly via migration from local refuge areas, where a reproductive and abundant meiofauna is thriving. These findings are important to consider in any potential future deep-sea mining scenario at deep-sea hydrothermal vents. Within our 4-year study period, we regularly observed vent habitats with tubeworm assemblages that became senescent and died, as vent fluid emissions locally stopped at patches within active vent sites. Senescent vents harbored a species rich mix of typical vent species as well as rare yet undescribed species. The senescent vents contributed significantly to diversity at the 9°N EPR with 55 macrofaunal species (11 singletons) and 74 meiofaunal species (19 singletons). Of these 129 species associated with senescent vents, 60 have not been reported from active vents. Tubeworms and other vent megafauna not only act as foundation species when alive but provide habitat also when dead, sustaining abundant and diverse small sized fauna.

Chapter 7 Pelagic Viruses, Bacteria and Archaea Markus Weinbauer, Markus WeinbauerSearch for more papers by this authorBranko Velimirov, Branko VelimirovSearch for more papers by this author Markus Weinbauer, Markus WeinbauerSearch for more papers by this authorBranko Velimirov, Branko VelimirovSearch for more papers by this author Book Editor(s):Christophe Migon, Christophe MigonSearch for more papers by this authorPaul Nival, Paul NivalSearch for more papers by this authorAntoine Sciandra, Antoine SciandraSearch for more papers by this author First published: 24 February 2020 https://doi.org/10.1002/9781119706960.ch7 AboutPDF ToolsRequest permissionExport citationAdd to favoritesTrack citation ShareShare Give accessShare full text accessShare full-text accessPlease review our Terms and Conditions of Use and check box below to share full-text version of article.I have read and accept the Wiley Online Library Terms and Conditions of UseShareable LinkUse the link below to share a full-text version of this article with your friends and colleagues. Learn more.Copy URL Share a linkShare onFacebookTwitterLinked InRedditWechat Summary Dissolved organic matter (DOM) is, for example, released or "exuded" from phytoplankton or produced by during feeding of zooplankton. Bacteria and archaea (together called prokaryotes) are the main users of DOM, and are consumed by flagellates and ciliates, which are eaten in turn by meso and macrozooplankton. In temperate marine ecosystems, the spring phytoplankton bloom is often a period during which changes in microbial activity are prominent. Prokaryotes and viruses are too small to be affected directly by turbulence. However, they can be influenced indirectly, e.g. when turbulence influences photosynthetic extracellular release or the formation of organic aggregates. Viral lysis and grazing by flagellates are the main factors causing mortality of prokaryotes, and thus shaping diversity and activity of prokaryotes. Data from experiment suggest that horizontal gene transfer could have a strong effect on the metabolic versatility and diversity of prokaryotes. The Mediterranean Sea in the Era of Global Change 1: 30 Years of Multidisciplinary Study of the Ligurian Sea RelatedInformation

Viral abundance in deep-sea environments is high. However, the biological, ecological and biogeochemical roles of viruses in the deep sea are under debate. In the present study, microcosm incubations of deep-sea bacterioplankton (2,000 m deep) with normal and reduced pressure of viral lysis were conducted in the western Pacific Ocean. We observed a negative effect of viruses on prokaryotic abundance, indicating the top-down control of bacterioplankton by virioplankton in the deep-sea. The decreased bacterial diversity and a different bacterial community structure with diluted viruses indicate that viruses are sustaining a diverse microbial community in deep-sea environments. Network analysis showed that relieving viral pressure decreased the complexity and clustering coefficients but increased the proportion of positive correlations for the potentially active bacterial community, which suggests that viruses impact deep-sea bacterioplankton interactions. Our study provides experimental evidences of the crucial role of viruses in microbial ecology and biogeochemistry in deep-sea ecosystems.

Recent research suggests that corals including cold-water corals harbor a diverse community of bacteria that are not only pathogens but also potential mutualists. Here we review data on bacterial community composition and diversity on the main cold-water corals framework builder species: Lophelia pertusa and Madrepora oculata. Sampling strategies such as box core, video grabs and remotely operated vehicle did not reveal strong differences between bacterial community composition as long as samples were used that looked 'not contaminated'. However, there were strong differences of bacterial diversity between the two coral species. An analysis of bacterial community composition by pyrosequencing of L. pertusa and M. oculata revealed for the Mediterranean Sea the presence of the potential mutualists already found in the Atlantic indicating a species-specific core microbiome. The data also suggest some biogeographical differences between the Mediterranean Sea and the North Atlantic for both coral species, however, this depends on the phylogenetic levels applied. In addition, there was also indication for a shared microbiome between the Mediterranean Sea and the Atlantic. Therefore species-specific bacterial associations seem to exist, whereas the biogeographical variability can be seen as adaptation to specific environmental conditions.