Plastics are pervasive in marine ecosystems and ubiquitous in both shallow and deep oceans. Microfibers, amongst other microscopic plastics, accumulate in deep sea sediments at concentrations up to four orders of magnitude higher than in surface waters. This is at odds with the fact that most microfibers are positively buoyant; therefore, it is hypothesized that settling aggregates are vectors for downward transport of microfibers in the ocean. However, little is known about the impact of microfibers on carbon export. We formed diatom aggregates with differing concentrations of microfibers using roller tanks and observed that microfiber addition stimulated aggregate formation, but decreased their structural cohesion and caused them to break apart more readily, resulting in smaller average sizes. Incorporation of positively buoyant microfibers into settling aggregates reduced their size-specific sinking velocities proportional to the microfiber concentration. Slower sinking may extend aggregate retention time in the upper ocean, thereby increasing the time available for organic matter remineralization in the upper water column. Here, we show that microfiber concentrations typical of those in the English Channel and Atlantic Ocean decrease potential export flux by 15-50%. Present day microfiber concentrations in surface waters may therefore be substantially reducing the efficiency of the biological carbon pump relative to the pre-plastic era.
The marine biological carbon pump is mainly driven by the interplay between aggregate sinking velocity and remineralization. Sinking velocity of natural marine aggregates is not routinely measured but often calculated using Stokes’ law, which does not consider size-dependent changes in porosity. We analyzed the flow fields around 81 in situ -formed aggregates using Particle Image Velocimetry (PIV) to determine the factors controlling aggregate settling. Using an independently derived scaling of porosity with size, we predicted the sinking velocity of laboratory-formed and in situ -formed aggregates with known densities. Small aggregates (<500 µm) have relatively lower porosities than large aggregates, and their increased compactness and density leads to higher size-specific settling velocities, and generally higher carbon-to-volume ratios. Applying our scaling approach to a global data set of vertical aggregate abundance and size distribution, we found that small aggregates contribute 40-70% to total carbon fluxes in situ . Teaser Improved sinking velocity prediction for marine aggregates highlights the contribution of small aggregates to carbon sequestration.
Next to iron (Fe), recent phytoplankton-enrichment experiments identified manganese (Mn) to (co-)limit Southern Ocean phytoplankton biomass and species composition. Since taxonomic diversity affects aggregation time and sinking rate, the efficiency of the biological carbon pump is directly affected by community structure. However, the impact of FeMn co-limitation on Antarctic primary production, community composition, and the subsequent export of carbon to depth requires more investigation. In situ samplings of 6 stations in the understudied southern Weddell Sea revealed that surface Fe and Mn concentrations, primary production, and carbon export rates were all low, suggesting a FeMn co-limited phytoplankton community. An Fe and Mn addition experiment examined how changes in the species composition drive the aggregation capability of a natural phytoplankton community. Primary production rates were highest when Fe and Mn were added together, due to an increased abundance of the colonial prymnesiophyte Phaeocystis antarctica. Although the community remained diatom dominated, the increase in Phaeocystis abundance led to highly carbon-enriched aggregates and a 4-fold increase in the carbon export potential compared to the control, whereas it only doubled in the Fe treatment. Based on the outcome of the FeMn-enrichment experiment, this region may suffer from FeMn co-limitation. As the Weddell Sea represents one of the most productive Antarctic marginal ice zones, our findings highlight that in response to greater Fe and Mn supply, changes in plankton community composition and primary production can have a disproportionally larger effect on the carbon export potential.
The marine biological carbon pump is mainly driven by the interplay between aggregate sinking velocity and remineralization. Sinking velocity of natural marine aggregates is not routinely measured but often calculated using Stokes’ law, which does not consider size-dependent changes in porosity. We analyzed the flow fields around 81 in situ -formed aggregates using Particle Image Velocimetry (PIV) to determine the factors controlling aggregate settling. Using an independently derived scaling of porosity with size, we predicted the sinking velocity of laboratory-formed and in situ -formed aggregates with known densities. Small aggregates (<500 µm) have relatively lower porosities than large aggregates, and their increased compactness and density leads to higher size-specific settling velocities, and generally higher carbon-to-volume ratios. Applying our scaling approach to a global data set of vertical aggregate abundance and size distribution, we found that small aggregates contribute 40-70% to total carbon fluxes in situ .Teaser Improved sinking velocity prediction for marine aggregates highlights the contribution of small aggregates to carbon sequestration.### Competing Interest StatementThe authors have declared no competing interest.
Phytoplankton forms the base of aquatic food webs and element cycling in diverse aquatic systems. The fate of phytoplankton-derived organic matter, however, often remains unresolved as it is controlled by complex, interlinked remineralization and sedimentation processes. We here investigate a rarely considered control mechanism on sinking organic matter fluxes: fungal parasites infecting phytoplankton. We demonstrate that bacterial colonization is promoted 3.5-fold on fungal-infected phytoplankton cells in comparison to non-infected cells in a cultured model pathosystem (diatom Synedra, fungal microparasite Zygophlyctis, and co-growing bacteria), and even ≥17-fold in field-sampled populations (Planktothrix, Synedra, and Fragilaria). Additional data obtained using the Synedra-Zygophlyctis model system reveals that fungal infections reduce the formation of aggregates. Moreover, carbon respiration is 2-fold higher and settling velocities are 11-48% lower for similar-sized fungal-infected vs. non-infected aggregates. Our data imply that parasites can effectively control the fate of phytoplankton-derived organic matter on a single-cell to single-aggregate scale, potentially enhancing remineralization and reducing sedimentation in freshwater and coastal systems.
A breeding colony of notothenioid icefish (Neopagetopsis ionah, Nybelin 1947) of globally unprecedented extent has been discovered in the southern Weddell Sea, Antarctica. The colony was estimated to cover at least similar to 240 km(2) of the eastern flank of the Filchner Trough, comprised of fish nests at a density of 0.26 nests per square meter, representing an estimated total of -60 million active nests and associated fish biomass of >60,000 tonnes. The majority of nests were each occupied by 1 adult fish guarding 1,735 eggs (+/- 433 SD). Bottom water temperatures measured across the nesting colony were up to 2 degrees C warmer than the surrounding bottom waters, indicating a spatial correlation between the modified Warm Deep Water (mWDW) upflow onto the Weddell Shelf and the active nesting area. Historical and concurrently collected seal movement data indicate that this concentrated fish biomass may be utilized by predators such as Weddell seals (Leptonychotes weddellii, Lesson 1826). Numerous degraded fish carcasses within and near the nesting colony suggest that, in death as well as life, these fish provide input for local food webs and influence local biogeochemical processing. To our knowledge, the area surveyed harbors the most spatially expansive continuous fish breeding colony discovered to date globally at any depth, as well as an exceptionally high Antarctic seafloor biomass. This discovery provides support for the establishment of a regional marine protected area in the Southern Ocean under the Convention on the Conservation of Antarctic Marine Living Resources (CCAMLR) umbrella.
Krill and salps are important for carbon flux in the Southern Ocean, but the extent of their contribution and the consequences of shifts in dominance from krill to salps remain unclear. We present a direct comparison of the contribution of krill and salp faecal pellets (FP) to vertical carbon flux at the Antarctic Peninsula using a combination of sediment traps, FP production, carbon content, microbial degradation, and krill and salp abundances. Salps produce 4-fold more FP carbon than krill, but the FP from both species contribute equally to the carbon flux at 300 m, accounting for 75% of total carbon. Krill FP are exported to 72% to 300 m, while 80% of salp FP are retained in the mixed layer due to fragmentation. Thus, declining krill abundances could lead to decreased carbon flux, indicating that the Antarctic Peninsula could become a less efficient carbon sink for anthropogenic CO 2 in future.
Over the past decades, two key grazers in the Southern Ocean (SO), krill and salps, have experienced drastic changes in their distribution and abundance, leading to increasing overlap of their habitats. Both species occupy different ecological niches and long-term shifts in their distributions are expected to have cascading effects on the SO ecosystem. However, studies directly comparing krill and salps are lacking. Here, we provide a direct comparison of the diet and fecal pellet composition of krill and salps using 18S metabarcoding and fatty acid markers. Neither species' diet reflected the composition of the plankton community, suggesting that in contrast to the accepted paradigm, not only krill but also salps are selective feeders. Moreover, we found that krill and salps had broadly similar diets, potentially enhancing the competition between both species. This could be augmented by salps' ability to rapidly reproduce in favorable conditions, posing further risks to krill populations.
Marine snow aggregates are microhabitats for diverse microbial communities with various active metabolic pathways. Rapid recycling and symbiotic transfer of nutrients within aggregates poses a significant challenge for accurately assessing aggregate‐associated turnover rates. Although single‐cell uptake measurements are well‐established for free‐living microorganisms, suitable methods for cells embedded in marine snow are currently lacking. Comparable cell‐specific measurements within sinking pelagic aggregates would have the potential to address core questions regarding aggregate‐associated fluxes. However, the capacity to perform microscale studies is limited by the difficulty of sampling and preserving the fragile aggregate structure. Furthermore, the application of nano‐scale secondary ion mass spectrometry (NanoSIMS) to aggregates is complicated by technical requirements related to vacuum and ablation resistance. Here, we present a NanoSIMS‐optimized method for fixation, embedding, and sectioning of marine snow. Stable isotope labeling of laboratory‐generated aggregates enabled visualization of label incorporation into prokaryotic and eukaryotic cells embedded in the aggregate structure. The current method is also amenable to various staining procedures, including transparent exopolymer particles, Coomassie stainable particles, nucleic acids, and eukaryotic cytoplasm. We demonstrate the potential for using structural stains to generate three‐dimensional (3D) models of marine snow and present a simplified calculation of porosity and fractal dimension. This multipurpose method enables combined investigations of 3D aggregate structure, spatial microbial distribution, and single‐cell activity within individual aggregates and provides new possibilities for future studies on microbial interactions and elemental uptake within marine snow.
The biological carbon pump is largely driven by the formation and sinking of marine snow. Because of their high organic matter content, marine snow aggregates are hotspots for microbial activity, and microbial organic matter degradation plays an important role in the attenuation of carbon fluxes to the deep sea. Our inability to examine and characterize microscale distributions of compounds making up the aggregate matrix, and of possible niches inside marine snow, has hindered our understanding of the basic processes governing marine carbon export and sequestration. To address this issue, we have adapted soft-embedding and sectioning to study the spatial structure and components of marine aggregates at high resolution. Soft-embedding enables rapid quantitative sampling of undisturbed marine aggregates from the water column and from sediment traps, followed by spatially resolved staining and characterization of substrates of the aggregate matrix and the microorganisms attached to it. Particular strengths of the method include in situ embedding in sediment traps and successful fluorescence in situ hybridization (FISH)-probe labeling, supporting studies of microbial diversity and ecology. The high spatial resolution achieved by thin-sectioning of soft-embedded aggregates offers the possibility for improved understanding of the composition and structure of marine snow, which directly influence settling velocity, microbial colonization and diversity, degradation rates, and carbon content. Our method will help to elucidate the small-scale processes underlying large-scale carbon cycling in the marine environment, which is especially relevant in the context of rising anthropogenic CO2 emissions and global change.
In the Arctic Ocean, climate change effects such as warming and ocean acidification (OA) are manifesting faster than in other regions. Yet, we are lacking a mechanistic understanding of the interactive effects of these drivers on Arctic primary producers. In the current study, one of the most abundant species of the Arctic Ocean, the prasinophyte Micromonas pusilla, was exposed to a range of different pCO(2) levels at two temperatures representing realistic current and future scenarios for nutrient-replete conditions. We observed that warming and OA synergistically increased growth rates at intermediate to high pCO(2) levels. Furthermore, elevated temperatures shifted the pCO(2) optimum of biomass production to higher levels. Based on changes in cellular composition and photophysiology, we hypothesise that the observed synergies can be explained by beneficial effects of warming on carbon fixation in combination with facilitated carbon acquisition under OA. Our findings help to understand the higher abundances of picoeukaryotes such as M. pusilla under OA, as has been observed in many mesocosm studies.
A previous study of 76 plant species on Spitsbergen in the High Arctic concluded that structures resembling arbuscular mycorrhizas were absent from roots. Here, we report a survey examining the roots of 13 grass and forb species collected from 12 sites on the island for arbuscular mycorrhizal (AM) colonisation. Of the 102 individuals collected, we recorded AM endophytes in the roots of 41 plants of 11 species (Alopecurus ovatus, Deschampsia alpina, Festuca rubra ssp. richardsonii, putative viviparous hybrids of Poa arctica and Poa pratensis, Poa arctica ssp. arctica, Trisetum spicatum, Coptidium spitsbergense, Ranunculus nivalis, Ranunculus pygmaeus, Ranunculus sulphureus and Taraxacum arcticum) sampled from 10 sites. Both coarse AM endophyte, with hyphae of 5–10 μm width, vesicles and occasional arbuscules, and fine endophyte, consisting of hyphae of 1–3 μm width and sparse arbuscules, were recorded in roots. Coarse AM hyphae, vesicles, arbuscules and fine endophyte hyphae occupied 1.0–30.7, 0.8–18.3, 0.7–11.9 and 0.7–12.8% of the root lengths of colonised plants, respectively. Principal component analysis indicated no associations between the abundances of AM structures in roots and edaphic factors. We conclude that the AM symbiosis is present in grass and forb roots on Spitsbergen.
Most studies of aquatic plankton focus on either macroscopic or microbial communities, and on either eukaryotes or prokaryotes. This separation is primarily for methodological reasons, but can overlook potential interactions among groups. Here we tested whether DNA metabarcoding of unfractionated water samples with universal primers could be used to qualitatively and quantitatively study the temporal dynamics of the total plankton community in a shallow temperate lake. Significant changes in the relative proportions of normalized sequence reads of eukaryotic and prokaryotic plankton communities over a 3-month period in spring were found. Patterns followed the same trend as plankton estimates measured using traditional microscopic methods. The bloom of a conditionally rare bacterial taxon belonging to Arcicella was characterized, which rapidly came to dominate the whole lake ecosystem and would have remained unnoticed without metabarcoding. The data demonstrate the potential of universal DNA metabarcoding applied to unfractionated samples for providing a more holistic view of plankton communities.
Most studies of biodiversity focus on either macroscopic or microbial communities, with little or no simultaneous study of eukaryotes and prokaryotes. We tested whether a universal metabarcoding approach could be used to study the total diversity and temporal dynamics of aquatic pico- to mesoplankton communities in a shallow temperate lake. The approach revealed significant changes in the relative abundance of eukaryotic and prokaryotic plankton communities over a period of three months. These patterns, based on sequencing reads, fit with counts using traditional methods. We also witnessed the bloom of a conditionally rare bacterial taxon belonging to Arcicella, a genus that has been largely overlooked in freshwaters. Our data demonstrate the potential of universal metabarcoding as a complement to traditional studies of plankton communities, and for long-term monitoring across a broad range of organisms.
We review the literature and find 16 studies from across Africa's savannas and woodlands where woody encroachment dominates. These small-scale studies are supplemented by an analysis of long-term continent-wide satellite data, specifically the Normalized Difference Vegetation Index (NDVI) time series from the Global Inventory Modeling and Mapping Studies (GIMMS) dataset. Using dry-season data to separate the tree and grass signals, we find 4.0% of non-rainforest woody vegetation in sub-Saharan Africa (excluding West Africa) significantly increased in NDVI from 1982 to 2006, whereas 3.52% decreased. The increases in NDVI were found predominantly to the north of the Congo Basin, with decreases concentrated in the Miombo woodland belt. We hypothesize that areas of increasing dry-season NDVI are undergoing woody encroachment, but the coarse resolution of the study and uncertain relationship between NDVI and woody cover mean that the results should be interpreted with caution; certainly, these results do not contradict studies finding widespread deforestation throughout the continent. However, woody encroachment could be widespread, and warrants further investigation as it has important consequences for the global carbon cycle and land-climate interactions.