Description: We estimate the global rate of biogenic silica production in the ocean to be between 200 and 280 × 1012 mol Si yr−1. The upper limit is derived from information on the primary productivity of the oceans, the relative contribution of diatoms to primary production and diatom Si/C ratios. The lower limit is derived independently using a multi‐compartment model of nutrient transport and biogenic particle flux, and field data on the balance between silica production and dissolution in the upper ocean. Our upper limit is 30–50% lower than several previous estimates, due to new data indicating lower values for both the relative contribution of diatoms to primary productivity and their Si/C ratios. Globally, at least 50% of the silica produced by diatoms in the euphotic zone dissolves in the upper 100 m, resulting in an estimated export of 100–140 × 1012 mol Si yr−l to the deep ocean. Our estimates correspond to a global mean rate of biogenic silica production between 0.6 and 0.8 mol Si m−2 yr−1. Incubation experiments indicate that silica production rates exceed that mean by a factor of 3–12 in coastal areas and are 2–4 times less than the global average in the oligotrophic mid‐ocean gyres. The mean silica production rate in waters overlying diatomaceous sediments (approximately 10–12% of the surface area of the oceans) is 0.7–1.2 mol Si m−2 yr−1. That rate is only slightly higher than the global average, indicating that the silica produced in those regions is only 10–25% of the global total. The estimated production of biogenic silica in surface waters of the mid‐ocean gyres is approximately equal to that for all major areas of opal sediment accumulation combined. Regional comparison of silica production and accumulation rates suggests a strongly bimodal character in the efficiency of opal preservation in the sea. In waters overlying diatom‐rich sediments 15–25% of the silica produced in the surface layer accumulates in the seabed, while virtually none of the silica produced in other areas is preserved. The global burial/production ratio of ˜ 3% is a composite of those two very different systems. The mechanisms leading to more efficient opal preservation in regions of silica accumulation are presently unknown, but they have no simple relationship to primary productivity. Regional differences in opal preservation appear to be controlled by factors such as low surface temperature, selective grazing and aggregate formation, which diminish the rate of silica dissolution in surface waters and/or accelerate its transport to the seafloor.

Description: Thriving in both epipelagic and mesopelagic layers, Rhizaria are biomineralizing protists, mixotrophs or flux-feeders, often reaching gigantic sizes. In situ imaging showed their contribution to oceanic carbon stock, but left their contribution to element cycling unquantified. Here, we compile a global dataset of 167,551 Underwater Vision Profiler 5 Rhizaria images, and apply machine learning models to predict their organic carbon and biogenic silica biomasses in the uppermost 1000 m. We estimate that Rhizaria represent up to 1.7% of mesozooplankton carbon biomass in the top 500 m. Rhizaria biomass, dominated by Phaeodaria, is more than twice as high in the mesopelagic than in the epipelagic layer. Globally, the carbon demand of mesopelagic, flux-feeding Phaeodaria reaches 0.46 Pg C y −1 , representing 3.8 to 9.2% of gravitational carbon export. Furthermore, we show that Rhizaria are a unique source of biogenic silica production in the mesopelagic layer, where no other silicifiers are present. Our global census further highlights the importance of Rhizaria for ocean biogeochemistry.

Description: Due to the major role played by diatoms in the biological pump of CO2, and to the presence of silica-rich sediments in areas that play a major role in air-sea CO2 exchange (e.g. the Southern Ocean and the Equatorial Pacific), opal has a strong potential as a proxy for paleoproductivity reconstructions. However, because of spatial variations in the biogenic silica preservation, and in the degree of coupling between the marine Si and C biogeochemical cycles, paleoreconstructions are not straitghtforward. A better calibration of this proxy in the modern ocean is required, which needs a good understanding of the mechanisms that control the Si cycle, in close relation to the carbon cycle. This review of the Si cycle in the modern ocean starts with the mechanisms that control the uptake of silicic acid (Si(OH)4) by diatoms and the subsequent silicification processes, the regulatory mechanisms of which are uncoupled. This has strong implications for the direct measurement in the field of the kinetics of Si(OH)4 uptake and diatom growth. It also strongly influences the Si:C ratio within diatoms, clearly linked to environmental conditions. Diatoms tend to dominate new production at marine ergoclines. At depth, they also succeed to form mats, which sedimentation is at the origin of laminated sediments and marine sapropels. The concentration of Si(OH)4 with respect to other macronutrients exerts a major influence on diatom dominance and on the rain ratio between siliceous and calcareous material, which severely impacts surface waters pCO2. A compilation of biogenic fluxes collected at about 40 sites by means of sediment traps also shows a remarkable pattern of increasing BSi:Corg ratio along the path of the "conveyor belt", accompanying the relative enrichment of waters in Si compared to N and P. This observation suggests an extension of the Si pump model described by Dugdale and Wilkerson (1989, doi:10.1038/34630), giving to Si(OH)4 a major role in the control of the rain ratio, which is of major importance in the global carbon cycle. The fate of the BSi produced in surface waters is then described, in relation to Corg, in terms of both dissolution and preservation mechanisms. Difficulties in quantifying the dissolution of biogenic silica in the water column as well as the sinking rates and forms of BSi to the deep, provide evidence for a major gap in our understanding of the mechanisms controlling the competition between retention in and export from surface waters. The relative influences of environmental conditions, seasonality, food web structure or aggregation are however explored. Quantitatively, assuming steady state, the measurements of the opal rain rate by means of sediment traps matches reasonably well those obtained by adding the recycling and burial fluxes in the underlying abyssal sediments, for most of the sites where such a comparison is possible. The major exception is the Southern Ocean where sediment focusing precludes the closing of mass balances. Focusing in fact is also an important aspect of the downward revision of the importance of Southern Ocean sediments in the global biogenic silica accumulation. Qualitatively, little is known about the duration of the transfer through the deep and the quality of the material that reaches the seabed, which is suggested to represent a major gap in our understanding of the processes governing the early diagenesis of BSi in sediments. The sediment composition (special emphasis on Al availability), the sedimentation rate or bioturbation are shown to exert an important control on the competition between dissolution and preservation of BSi in sediments. It is suggested that a primary control on the kinetic and thermodynamic properties of BSi dissolution, both in coastal and abyssal sediments, is exerted by water column processes, either occuring in surface waters during the formation of the frustules, or linked to the transfer of the particles through the water column, which duration may influence the quality of the biogenic rain. This highlights the importance of studying the factors controlling the degree of coupling between pelagic and benthic processes in various regions of the world ocean, and its consequences, not only in terms of benthic biology but also for the constitution of the sediment archive. The last section, first calls for the end of the "NPZD" models, and for the introduction of processes linked to the Si cycle, into models describing the phytoplankton cycles in surface waters and the early diagenesis of BSi in sediments. It also calls for the creation of an integrated 1-D diagnostic model of the Si:C coupling, for a better understanding of the interactions between surface waters, deep waters and the upper sedimentary column. The importance of Si(OH)4 in the control of the rain ratio and the improved parametrization of the Si cycle in the 1-D diagnostic models should lead to a reasonable incorporation of the Si cycle into 3-D regional circulation models and OGCMs, with important implications for climate change studies and paleoreconstructions at regional and global scale.

Description: During the EBENE cruise (November 1996), distributions of biogenic silica concentration and production rates were investigated in the surface waters of the equatorial Pacific (180°W, from 8°S to 8°N), with particular emphasis on the limitation of the biogenic silica production by ambient silicic acid concentrations. Integrated over the depth of the euphotic layer, concentrations of biogenic silica and production rates were maximum at the Equator (8.0 and 2.6 mmol/m**2/d) and decreased more or less symmetrically polewards. Contribution of diatoms to the new production was estimated indirectly, comparing biogenic silica production rates and available data of new and export production in the same area. This comparison shows that new production in the equatorial area could mostly be sustained by diatoms, accounting for the major part of the exported flux of organic carbon. Kinetics experiments of silicic acid enrichment were performed. Half saturation constants were 1.57 µM at 3°S and 2.42 µM at the Equator close to the ambient concentrations. The corresponding Vmax values for Si uptake were 0.028/h at 3°S and 0.052/h at the equator. Experiments also show that in situ rates were restricted to 13-78% of Vmax, depending on ambient silicic acid concentrations. This work provides the first direct evidence that the rate of Si uptake by diatom populations of the equatorial Pacific is limited by the ambient concentration of silicic acid. However, such Si limitation might not be sufficient in itself to explain the low diatom growth rates observed, and additional limitation is suggested. One hypothesis that is consistent with the results of Fe limitation studies is that Fe and Si limitations may interact, rather than just being a mutually exclusive explanation for the HNLC character of the system.

Description: © The Author(s), 2021. This article is distributed under the terms of the Creative Commons Attribution License. The definitive version was published in Treguer, P. J., Sutton, J. N., Brzezinski, M., Charette, M. A., Devries, T., Dutkiewicz, S., Ehlert, C., Hawkings, J., Leynaert, A., Liu, S. M., Monferrer, N. L., Lopez-Acosta, M., Maldonado, M., Rahman, S., Ran, L., & Rouxel, O. Reviews and syntheses: the biogeochemical cycle of silicon in the modern ocean. Biogeosciences, 18(4), (2021): 1269-1289, https://doi.org/10.5194/bg-18-1269-2021.

Description: The element silicon (Si) is required for the growth of silicified organisms in marine environments, such as diatoms. These organisms consume vast amounts of Si together with N, P, and C, connecting the biogeochemical cycles of these elements. Thus, understanding the Si cycle in the ocean is critical for understanding wider issues such as carbon sequestration by the ocean's biological pump. In this review, we show that recent advances in process studies indicate that total Si inputs and outputs, to and from the world ocean, are 57 % and 37 % higher, respectively, than previous estimates. We also update the total ocean silicic acid inventory value, which is about 24 % higher than previously estimated. These changes are significant, modifying factors such as the geochemical residence time of Si, which is now about 8000 years, 2 times faster than previously assumed. In addition, we present an updated value of the global annual pelagic biogenic silica production (255 Tmol Si yr−1) based on new data from 49 field studies and 18 model outputs, and we provide a first estimate of the global annual benthic biogenic silica production due to sponges (6 Tmol Si yr−1). Given these important modifications, we hypothesize that the modern ocean Si cycle is at approximately steady state with inputs =14.8(±2.6) Tmol Si yr−1 and outputs =15.6(±2.4) Tmol Si yr−1. Potential impacts of global change on the marine Si cycle are discussed.

Description: This work was supported by the French National Research Agency (18-CEO1-0011-01) and by the Spanish Ministry of Science, Innovation and Universities (PID2019-108627RB-I00).

Description: Thriving in both epipelagic and mesopelagic layers, Rhizaria are biomineralizing protists, mixotrophs or flux-feeders, often reaching gigantic sizes. In situ imaging showed their contribution to oceanic carbon stock, but left their contribution to element cycling unquantified. Here, we compile a global dataset of 167,551 Underwater Vision Profiler 5 Rhizaria images, and apply machine learning models to predict their organic carbon and biogenic silica biomasses in the uppermost 1000 m. We estimate that Rhizaria represent up to 1.7% of mesozooplankton carbon biomass in the top 500 m. Rhizaria biomass, dominated by Phaeodaria, is more than twice as high in the mesopelagic than in the epipelagic layer. Globally, the carbon demand of mesopelagic, flux-feeding Phaeodaria reaches 0.46 Pg C y〈jats:sup〉−1〈/jats:sup〉, representing 3.8 to 9.2% of gravitational carbon export. Furthermore, we show that Rhizaria are a unique source of biogenic silica production in the mesopelagic layer, where no other silicifiers are present. Our global census further highlights the importance of Rhizaria for ocean biogeochemistry.