Elongation enhances encounter rates between phytoplankton in turbulence.

Phytoplankton come in a stunning variety of shapes but elongated morphologies dominate-typically 50% of species have aspect ratio above 5, and bloom-forming species often form chains whose aspect ratios can exceed 100. How elongation affects encounter rates between phytoplankton in turbulence has remained unknown, yet encounters control the formation of marine snow in the ocean. Here, we present simulations of encounters among elongated phytoplankton in turbulence, showing that encounter rates between neutrally buoyant elongated cells are up to 10-fold higher than for spherical cells and even higher when cells sink. Consequently, we predict that elongation can significantly speed up the formation of marine snow compared to spherical cells. This unexpectedly large effect of morphology in driving encounter rates among plankton provides a potential mechanistic explanation for the rapid clearance of many phytoplankton blooms.

Vertical Flux of Microplastics in the Deep Subtropical Pacific Ocean: Moored Sediment-Trap Observations within the Kuroshio Extension Recirculation Gyre.

The Kuroshio Extension recirculation gyre in the western North Pacific is an accumulation site of plastic litter transported by the Kuroshio Current. A sediment trap was moored at a depth of 4900 m at Station KEO within the Kuroshio Extension recirculation gyre, and the vertical flux of microplastics in sinking particles of size <1 mm was observed. Forty-one sediment-trap samples collected from July 1, 2014, to October 2, 2016, were analyzed with a micro-Fourier transform infrared spectrometer and microplastics were detected in all samples. Seventeen polymer types were identified, and 90% of the microplastics were less than 100 µm in size. Microplastic sinking was driven by the action of the biological pump, which was in turn driven by seasonal variations in solar radiation and increased surface primary production typical of the spring season. Microplastic mass flux varied from 4.5 × 10-3 to 0.38 mg m-2 day-1 during the sampling period, with a mean and standard deviation of 0.054 ± 0.075 mg m-2 day-1. Extrapolating the annual microplastic mass flux at Station KEO to the entire Kuroshio Extension recirculation gyre, it is estimated that 0.028 million metric tons of microplastics are transported annually to 4900 m depth in this area.

A Multi-Stage Progressive Network with Feature Transmission and Fusion for Marine Snow Removal.

Improving underwater image quality is crucial for marine detection applications. However, in the marine environment, captured images are often affected by various degradation factors due to the complexity of underwater conditions. In addition to common color distortions, marine snow noise in underwater images is also a significant issue. The backscatter of artificial light on marine snow generates specks in images, thereby affecting image quality, scene perception, and subsequently impacting downstream tasks such as target detection and segmentation. Addressing the issues caused by marine snow noise, we have designed a new network structure. In this work, a novel skip-connection structure called a dual channel multi-scale feature transmitter (DCMFT) is implemented to reduce information loss during downsampling in the feature encoding and decoding section. Additionally, in the feature transfer process for each stage, iterative attentional feature fusion (iAFF) modules are inserted to fully utilize marine snow features extracted at different stages. Finally, to further optimize the network's performance, we incorporate the multi-scale structural similarity index (MS-SSIM) into the loss function to ensure more effective convergence during training. Through experiments conducted on the Marine Snow Removal Benchmark (MSRB) dataset with an augmented sample size, our method has achieved significant results. The experimental results demonstrate that our approach excels in removing marine snow noise, with a peak signal-to-noise ratio reaching 38.9251 dB, significantly outperforming existing methods.

On the collision of rods in a quiescent fluid.

Rods settling under gravity in a quiescent fluid can overcome the bottleneck associated with aggregation of equal-size spheres because they collide by virtue of their orientation-dependent settling velocity. We find the corresponding collision kernel [Formula: see text], where l, A, and [Formula: see text] are the rods' length, aspect ratio (length divided by width), and volume, respectively, [Formula: see text] is the density difference between rods and fluid, µ is the fluid's dynamic viscosity, g is the gravitational acceleration, and [Formula: see text] is a geometrical parameter. We apply this formula to marine snow formation following a phytoplankton bloom. Over a broad range of aspect ratios, the formula predicts a similar or higher encounter rate between rods as compared to the encounter rate between (equal volume) spheres aggregating either by differential settling or due to turbulence. Since many phytoplankton species are elongated, these results suggest that collisions induced by the orientation-dependent settling velocity can contribute significantly to marine snow formation, and that marine snow composed of elongated phytoplankton cells can form at high rates also in the absence of turbulence.

Effect of Nano-Silver on Formation of Marine Snow and the Underlying Microbial Mechanism.

Roller experiments were conducted to explore the effect of nano-silver on the formation of marine snow and the underlying microbial mechanism. With the increasing concentration of nano-Ag from 1 ng/L to 1 mg/L, the formation and aggregation of marine snow particles were solidly suppressed in a dose-dependent pattern. Moreover, the formed marine snows tended to be thinner fibrous particles with smaller size and increased edge smoothness and compactness in the presence of nano-Ag. The microbial analyses indicated that nano-Ag not only inhibited the development of biomass but also changed the species composition and functional profile of the microbial community. Nano-Ag obviously inhibited most of the abundant species, except for some myxobacteria, which is unfavorable for the microbial community stability. For the microbial functions, some major biological processes including the growth, metabolic, and cellular processes were also inhibited by the high dosage of nano-Ag. The strong microbial inhibition of nano-Ag would contribute to the suppression on the formation of marine snow. Specifically, the function genes of extracellular polymeric substance synthesis and secretion were significantly reduced by nano-Ag, which might be the key and straight microbial factor in suppressing the formation of marine snow.

Hitchhiking into the Deep: How Microplastic Particles are Exported through the Biological Carbon Pump in the North Atlantic Ocean.

Understanding residence times of plastic in the ocean is a major knowledge gap in plastic pollution studies. Observations report a large mismatch between plastic load estimates from worldwide production and disposal and actual plastics floating at the sea surface. Surveys of the water column, from the surface to the deep sea, are rare. Most recent work, therefore, addressed the "missing plastic" question using modeling or laboratory approaches proposing biofouling and degradation as the main removal processes in the ocean. Through organic matrices, plastic can affect the biogeochemical and microbial cycling of carbon and nutrients. For the first time, we provide in situ measured vertical fluxes of microplastics deploying drifting sediment traps in the North Atlantic Gyre from 50 m down to 600 m depth, showing that through biogenic polymers plastic can be embedded into rapidly sinking particles also known as marine snow. We furthermore show that the carbon contained in plastic can represent up to 3.8% of the total downward flux of particulate organic carbon. Our results shed light on important pathways regulating the transport of microplastics in marine systems and on potential interactions with the marine carbon cycle, suggesting microplastic removal through the "biological plastic pump".

Using High-Sensitivity Lipidomics To Assess Microscale Heterogeneity in Oceanic Sinking Particles and Single Phytoplankton Cells.

Sinking particulate organic matter (POM) is a primary component of the ocean's biological carbon pump that is responsible for carbon export from the surface to the deep sea. Lipids derived from plankton comprise a significant fraction of sinking POM. Our understanding of planktonic lipid biosynthesis and the subsequent degradation of lipids in sinking POM is based on the analysis of bulk samples that combine many millions of plankton cells or dozens of sinking particles, which averages out natural heterogeneity. We developed and applied a nanoflow high-performance liquid-chromatography electrospray-ionization high-resolution accurate-mass mass spectrometry lipidomic method to show that two types of sinking particles─marine snow and fecal pellets─collected in the western North Atlantic Ocean have distinct lipidomes, providing new insights into their sources and degradation that would not be apparent from bulk samples. We pressed the limit of this approach by examining individual diatom cells from a single culture, finding marked lipid heterogeneity, possibly indicative of fundamental mechanisms underlying cell division. These single-cell data confirm that even cultures of phytoplankton cells should be viewed as mixtures of physiologically distinct populations. Overall, this work reveals previously hidden lipidomic heterogeneity in natural POM and phytoplankton cells, which may provide critical new insights into microscale chemical and microbial processes that control the export of sinking POM.

Vertical distribution of particle-associated and free-living ammonia-oxidizing archaea in Suruga Bay, a deep coastal embayment of Japan.

We analyzed the vertical distributions of ammonia-oxidizing archaea (AOA) in terms of abundance in Suruga Bay, Japan. We distinguished particle-associated (PA) from free-living (FL) assemblages. According to quantitative PCR measurements of the ammonia monooxygenase subunit A gene (amoA), most marine AOA were in an FL state. The vertical distributions of PA AOA ecotypes differed from the general trend; the Shallow Marine clade was dominant in both the surface and deep layers. Thus, although PA AOA account for a small percentage of AOA abundance, they have a community structure distinct from that of FL AOA in planktonic environments. Marine particles should be investigated further as an unexplored niche of AOA in the ocean.

Morphology, composition and aggregation mechanisms of soft bioflocs in marine snow and activated sludge: A comparative review.

Conditions that lead to marine snow formation and aggregates that constitute the marine snow have similarities with the soft bioflocs that form during wastewater treatment by activated sludge process. Analysis of the conditions and similarities of the soft bioflocs in these two aquatic environments provide insight for the processes that lead to formation and growth of hydrated aggregates consisting of both living and nonliving particles, their chemical and biolocial composition, settling/suspension behavior, and contributing factors for their structure and morphology. This literature review provides a comparative analysis of the soft aggregates that form in marine and wastewater environments to characterize the conditions for formation and growth of highly hydrated aggregates consisting of microorganisms, suspended solids and large molecules. The marine snow and bioflocs that form in wastewater are visually similar and even contain microorganisms that are of similar type (i.e., Zoogloea, filamentous bacteria). During wastewater treatment, the microorganisms are not stressed and exopolymeric substances (EPS) produced have shorter molecules and higher protein content while EPS produced by the marine organisms are significantly larger in molecular size (by orders of magnitude) and have higher carbohydrate content.

An examination of the factors influencing mercury and methylmercury particulate distributions, methylation and demethylation rates in laboratory-generated marine snow.

In the marine environment, settling particulates have been widely studied for their role as effective vertical transporters of nutrients and metals scavenged from the euphotic zone to the benthos. These particulates are composed of transparent exopolymers, plankton and bacterial cells, detritus and organic matter, and form various size fractions from colloids (<0.2µm) to aggregates, and finally marine snow (>300 µm). As marine snow forms in the water column, anoxic layers form around and within the aggregation potentially creating a prime environment for the methylation of mercury (Hg), which occurs primarily in low oxygen environments. To examine this process, marine aggregates were produced from sieved estuarine seawater (100 µm) in 1-L glass bottles spiked with stable isotope enriched methylmercury (CH3199Hg) and inorganic mercury (200Hg(II)) at 18° C using a roller-table. After the rolling period, different particle-size fractions were collected and analyzed, including: visible marine snow (>300µm), particulates 8 to 300 µm, and particulates 0.2 to 8µm. Particulate analysis indicated higher incorporation of both forms of Hg into marine snow compared to unrolled treatments, with greater incorporation of 200Hg(II) than CH3199Hg. In addition, inorganic Hg was methylated and CH3Hg was demethylated in the larger particulate fractions (>8µm). Methylation and demethylation rates were assessed based on changes in isotopic composition of Hg(II) and CH3Hg, and found to be comparable to methylation rates found in sediments. These results indicate that net Hg methylation can occur in marine snow and smaller aggregates in oxic coastal waters, and that this net formation of CH3Hg may be an important source of CH3Hg in both coastal and open ocean surface environments.

Marine bacteria Alteromonas spp. require UDP-glucose-4-epimerase for aggregation and production of sticky exopolymer.

The physiology and ecology of particle-associated marine bacteria are of growing interest, but our knowledge of their aggregation behavior and mechanisms controlling their association with particles remains limited. We have found that a particle-associated isolate, Alteromonas sp. ALT199 strain 4B03, and the related type-strain A. macleodii 27126 both form large (>500 µm) aggregates while growing in rich medium. A non-clumping variant (NCV) of 4B03 spontaneously arose in the lab, and whole-genome sequencing revealed a partial deletion in the gene encoding UDP-glucose-4-epimerase (galEΔ308-324). In 27126, a knock-out of galE (ΔgalE::kmr) resulted in a loss of aggregation, mimicking the NCV. Microscopic analysis shows that both 4B03 and 27126 rapidly form large aggregates, whereas their respective galE mutants remain primarily as single planktonic cells or clusters of a few cells. Strains 4B03 and 27126 also form aggregates with chitin particles, but their galE mutants do not. Alcian Blue staining shows that 4B03 and 27126 produce large transparent exopolymer particles (TEP), but their galE mutants are deficient in this regard. This study demonstrates the capabilities of cell-cell aggregation, aggregation of chitin particles, and production of TEP in strains of Alteromonas, a widespread particle-associated genus of heterotrophic marine bacteria. A genetic requirement for galE is evident for each of the above capabilities, expanding the known breadth of requirement for this gene in biofilm-related processes. IMPORTANCE: Heterotrophic marine bacteria have a central role in the global carbon cycle. Well-known for releasing CO2 by decomposition and respiration, they may also contribute to particulate organic matter (POM) aggregation, which can promote CO2 sequestration via the formation of marine snow. We find that two members of the prevalent particle-associated genus Alteromonas can form aggregates comprising cells alone or cells and chitin particles, indicating their ability to drive POM aggregation. In line with their multivalent aggregation capability, both strains produce TEP, an excreted polysaccharide central to POM aggregation in the ocean. We demonstrate a genetic requirement for galE in aggregation and large TEP formation, building our mechanistic understanding of these aggregative capabilities. These findings point toward a role for heterotrophic bacteria in POM aggregation in the ocean and support broader efforts to understand bacterial controls on the global carbon cycle based on microbial activities, community structure, and meta-omic profiling.

Identification of the bacterial community that degrades phenanthrene sorbed to polystyrene nanoplastics using DNA-based stable isotope probing.

In the Anthropocene, plastic pollution has become a new environmental biotope, the so-called plastisphere. In the oceans, nano- and micro-sized plastics are omnipresent and found in huge quantities throughout the water column and sediment, and their large surface area-to-volume ratio offers an excellent surface to which hydrophobic chemical pollutants (e.g. petrochemicals and POPs) can readily sorb to. Our understanding of the microbial communities that breakdown plastic-sorbed chemical pollutants, however, remains poor. Here, we investigated the formation of 500 nm and 1000 nm polystyrene (PS) agglomerations in natural seawater from a coastal environment, and we applied DNA-based stable isotope probing (DNA-SIP) with the 500 nm PS sorbed with isotopically-labelled phenanthrene to identify the bacterial members in the seawater community capable of degrading the hydrocarbon. Whilst we observed no significant impact of nanoplastic size on the microbial communities associated with agglomerates that formed in these experiments, these communities were, however, significantly different to those in the surrounding seawater. By DNA-SIP, we identified Arcobacteraceae, Brevundimonas, Comamonas, uncultured Comamonadaceae, Delftia, Sphingomonas and Staphylococcus, as well as the first member of the genera Acidiphilum and Pelomonas to degrade phenanthrene, and of the genera Aquabacterium, Paracoccus and Polymorphobacter to degrade a hydrocarbon. This work provides new information that feeds into our growing understanding on the fate of co-pollutants associated with nano- and microplastics in the ocean.

Comparing microbial communities in mucilage and seawater samples: Metagenomic insights into mucilage formation in the Marmara Sea.

Marine environments are subject to various naturally occurring phenomena, including marine snow and mucilage. In 2021, the rapid emergence of mucilage in the Marmara Sea raised concerns about its environmental impact. This study investigates the microbial communities in mucilage and seawater samples from the Marmara Sea using metagenomic-scale comparative analyses. The results indicate significant differences in microbial composition and diversity, with mucilage samples showing higher levels of polysaccharide biosynthesis-related enzymes. Over 50% of reads in mucilage samples remained unclassified (dark matter), highlighting unknown microbial taxa. Clean seawater was characterized by a higher presence of Euryarchaeota, Proteobacteria, and Rhodothermaeota, while Chlamydiae and Fusobacteria were dominant in mucilage. The study underscores the necessity for comprehensive metagenomic analyses to understand microbial roles in mucilage formation and persistence. Early detection of microbial shifts could serve as a warning system for mucilage outbreaks, aiding in the development of management strategies.

Marine particle size-fractionation indicates organic matter is processed by differing microbial communities on depth-specific particles.

Passive sinking flux of particulate organic matter in the ocean plays a central role in the biological carbon pump and carbon export to the ocean's interior. Particle-associated microbes colonize particulate organic matter, producing "hotspots" of microbial activity. We evaluated variation in particle-associated microbial communities to 500 m depth across four different particle size fractions (0.2-1.2, 1.2-5, 5-20, >20 µm) collected using in situ pumps at the Bermuda Atlantic Time-series Study site. In situ pump collections capture both sinking and suspended particles, complementing previous studies using sediment or gel traps, which capture only sinking particles. Additionally, the diagenetic state of size-fractionated particles was examined using isotopic signatures alongside microbial analysis. Our findings emphasize that different particle sizes contain distinctive microbial communities, and each size category experiences a similar degree of change in communities over depth, contradicting previous findings. The robust patterns observed in this study suggest that particle residence times may be long relative to microbial succession rates, indicating that many of the particles collected in this study may be slow sinking or neutrally buoyant. Alternatively, rapid community succession on sinking particles could explain the change between depths. Complementary isotopic analysis of particles revealed significant differences in composition between particles of different sizes and depths, indicative of organic particle transformation by microbial hydrolysis and metazoan grazing. Our results couple observed patterns in microbial communities with the diagenetic state of associated organic matter and highlight unique successional patterns in varying particle sizes across depth.

Porous marine snow differentially benefits chemotactic, motile, and nonmotile bacteria.

Particulate organic carbon settling through the marine water column is a key process that regulates the global climate by sequestering atmospheric carbon. The initial colonization of marine particles by heterotrophic bacteria represents the first step in recycling this carbon back to inorganic constituents-setting the magnitude of vertical carbon transport to the abyss. Here, we demonstrate experimentally using millifluidic devices that, although bacterial motility is essential for effective colonization of a particle leaking organic nutrients into the water column, chemotaxis specifically benefits at intermediate and higher settling velocities to navigate the particle boundary layer during the brief window of opportunity provided by a passing particle. We develop an individual-based model that simulates the encounter and attachment of bacterial cells with leaking marine particles to systematically evaluate the role of different parameters associated with bacterial run-and-tumble motility. We further use this model to explore the role of particle microstructure on the colonization efficiency of bacteria with different motility traits. We find that the porous microstructure facilitates additional colonization by chemotactic and motile bacteria, and fundamentally alters the way nonmotile cells interact with particles due to streamlines intersecting with the particle surface.

Active sinking particles: sessile suspension feeders significantly alter the flow and transport to sinking aggregates.

Sinking or sedimentation of biological aggregates plays a critical role in carbon sequestration in the ocean and in vertical material fluxes in wastewater treatment plants. In both these contexts, the sinking aggregates are 'active', since they are biological hot-spots and are densely colonized by microorganisms including bacteria and sessile protists, some of which generate feeding currents. However, the effect of these feeding currents on the sinking rates, trajectories and mass transfer to these 'active sinking particles' has not previously been studied. Here, we use a novel scale-free vertical tracking microscope (a.k.a. gravity machine; Krishnamurthy et al. 2020 Nat. Methods 17, 1040-1051 (doi:10.1038/s41592-020-0924-7)) to follow model sinking aggregates (agar spheres) with attached protists (Vorticella convallaria), sinking over long distances while simultaneously measuring local flows. We find that activity due to attached V. convallaria causes significant changes to the flow around aggregates in a dynamic manner and reshapes mass transport boundary layers. Further, we find that activity-mediated local flows along with sinking modify the encounter and plume cross-sections of the aggregate and induce sustained aggregate rotations. Overall, our work shows the important role of biological activity in shaping the near-field flows around aggregates with potentially important effects on aggregate fate and material fluxes.

Interaction of silver nanoparticles with marine/lake snow in early formation stage.

Marine and lake snows play an important ecological role in aquatic systems, and recent researches have also revealed their interactions with various pollutants. In this paper, the interaction of silver nanoparticles (Ag-NPs), a typical nano-pollutant, with marine/lake snow in the early formation stage was investigated by roller table experiments. Results indicated Ag-NPs promoted the accumulation of larger marine snow flocs while inhibited the development of lake snow. The promotion effect of AgNPs might be attributed to their oxidative dissolution into low-toxic silver chloride complexes in seawater, and the subsequent incorporation into marine snow, which would enhance the rigidity and strength of larger flocs and favor the development of biomass. Conversely, Ag-NPs mainly existed in the form of colloidal nanoparticles in lake water and their strong antimicrobial effect suppressed the growths of biomass and lake snow. In addition, Ag-NPs could also affect the microbial community of marine/lake snow, including impact on microbial diversity, and elevation on abundances of extracellular polymeric substances (EPS) synthesis genes and silver resistance genes. This work has deepened our understanding of the fate and ecological effect of Ag-NPs via the interaction with marine/lake snow in aquatic environments.

Carbon Export in the Ocean: A Biologist's Perspective.

Understanding the nature of organic matter flux in the ocean remains a major goal of oceanography because it impacts some of the most important processes in the ocean. Sinking particles are important for carbon dioxide removal from the atmosphere and its movement to the deep ocean. They also feed life below the ocean's productive surface and sustain life in the deep sea, in addition to depositing organic matter on the seafloor. However, the magnitude of all of these processes is dependent on the transformation of sinking particles during their journey through the water column. This review focuses on the movement of organic matter from the surface ocean to the deep sea via the biological carbon pump and examines the processes that prevent this downward movement-namely, attenuation via microbial colonization and zooplankton feeding. It also discusses how the depth-specific interactions among microbes, zooplankton, and aggregates determine carbon export as well as nutrient recycling, which in turn impact ocean production and Earth's climate.

Bacteria and Archaea Regulate Particulate Organic Matter Export in Suspended and Sinking Marine Particle Fractions.

The biological carbon pump (BCP) in the Southern Ocean is driven by phytoplankton productivity and is a significant organic matter sink. However, the role of particle-attached (PA) and free-living (FL) prokaryotes (bacteria and archaea) and their diversity in influencing the efficiency of the BCP is still unclear. To investigate this, we analyzed the metagenomes linked to suspended and sinking marine particles from the Sub-Antarctic Southern Ocean Time Series (SOTS) by deploying a Marine Snow Catcher (MSC), obtaining suspended and sinking particulate material, determining organic carbon and nitrogen flux, and constructing metagenome-assembled genomes (MAGs). The suspended and sinking particle-pools were dominated by bacteria with the potential to degrade organic carbon. Bacterial communities associated with the sinking fraction had more genes related to the degradation of complex organic carbon than those in the suspended fraction. Archaea had the potential to drive nitrogen metabolism via nitrite and ammonia oxidation, altering organic nitrogen concentration. The data revealed several pathways for chemoautotrophy and the secretion of recalcitrant dissolved organic carbon (RDOC) from CO2, with bacteria and archaea potentially sequestering particulate organic matter (POM) via the production of RDOC. These findings provide insights into the diversity and function of prokaryotes in suspended and sinking particles and their role in organic carbon/nitrogen export in the Southern Ocean. IMPORTANCE The biological carbon pump is crucial for the export of particulate organic matter in the ocean. Recent studies on marine microbes have shown the profound influence of bacteria and archaea as regulators of particulate organic matter export. Yet, despite the importance of the Southern Ocean as a carbon sink, we lack comparable insights regarding microbial contributions. This study provides the first insights regarding prokaryotic contributions to particulate organic matter export in the Southern Ocean. We reveal evidence that prokaryotic communities in suspended and sinking particle fractions harbor widespread genomic potential for mediating particulate organic matter export. The results substantially enhance our understanding of the role played by microorganisms in regulating particulate organic matter export in suspended and sinking marine fractions in the Southern Ocean.

Machine Learning for the Study of Plankton and Marine Snow from Images.

Quantitative imaging instruments produce a large number of images of plankton and marine snow, acquired in a controlled manner, from which the visual characteristics of individual objects and their in situ concentrations can be computed. To exploit this wealth of information, machine learning is necessary to automate tasks such as taxonomic classification. Through a review of the literature, we highlight the progress of those machine classifiers and what they can and still cannot be trusted for. Several examples showcase how the combination of quantitative imaging with machine learning has brought insights on pelagic ecology. They also highlight what is still missing and how images could be exploited further through trait-based approaches. In the future, we suggest deeper interactions with the computer sciences community, the adoption of data standards, and the more systematic sharing of databases to build a global community of pelagic image providers and users.