Cloud shadows drive vertical migrations of deep-dwelling marine life.

Many zooplankton and fishes vertically migrate on a diel cycle to avoid predation, moving from their daytime residence in darker, deep waters to prey-rich surface waters to feed at dusk and returning to depth before dawn. Vertical migrations also occur in response to other processes that modify local light intensity, such as storms, eclipses, and full moons. We observed rapid, high-frequency migrations, spanning up to 60 m, of a diel vertically migrating acoustic scattering layer with a daytime depth of 300 m in the subpolar Northeastern Pacific Ocean. The depth of the layer was significantly correlated, with an ∼5-min lag, to cloud-driven variability in surface photosynthetically available radiation. A model of isolume-following swimming behavior reproduces the observed layer depth and suggests that the high-frequency migration is a phototactic response to absolute light level. Overall, the cumulative distance traveled per day in response to clouds was at least 36% of the round-trip diel migration distance. This previously undescribed phenomenon has implications for the metabolic requirements of migrating animals while at depth and highlights the powerful evolutionary adaptation for visual predator avoidance.

The Outsized Role of Salps in Carbon Export in the Subarctic Northeast Pacific Ocean.

Periodic blooms of salps (pelagic tunicates) can result in high export of organic matter, leading to an "outsized" role in the ocean's biological carbon pump (BCP). However, due to their episodic and patchy nature, salp blooms often go undetected and are rarely included in measurements or models of the BCP. We quantified salp-mediated export processes in the northeast subarctic Pacific Ocean in summer of 2018 during a bloom of Salpa aspera. Salps migrated from 300 to 750 m during the day into the upper 100 m at night. Salp fecal pellet production comprised up to 82% of the particulate organic carbon (POC) produced as fecal pellets by the entire epipelagic zooplankton community. Rapid sinking velocities of salp pellets (400-1,200 m d-1) and low microbial respiration rates on pellets (<1% of pellet C respired day-1) led to high salp pellet POC export from the euphotic zone-up to 48% of total sinking POC across the 100 m depth horizon. Salp active transport of carbon by diel vertical migration and carbon export from sinking salp carcasses was usually <10% of the total sinking POC flux. Salp-mediated export markedly increased BCP efficiency, increasing by 1.5-fold the proportion of net primary production exported as POC across the base of the euphotic zone and by 2.6-fold the proportion of this POC flux persisting 100 m below the euphotic zone. Salps have unique and important effects on ocean biogeochemistry and, especially in low flux settings, can dramatically increase BCP efficiency and thus carbon sequestration.

A Visual Tour of Carbon Export by Sinking Particles.

To better quantify the ocean's biological carbon pump, we resolved the diversity of sinking particles that transport carbon into the ocean's interior, their contribution to carbon export, and their attenuation with depth. Sinking particles collected in sediment trap gel layers from four distinct ocean ecosystems were imaged, measured, and classified. The size and identity of particles was used to model their contribution to particulate organic carbon (POC) flux. Measured POC fluxes were reasonably predicted by particle images. Nine particle types were identified, and most of the compositional variability was driven by the relative contribution of aggregates, long cylindrical fecal pellets, and salp fecal pellets. While particle composition varied across locations and seasons, the entire range of compositions was measured at a single well-observed location in the subarctic North Pacific over one month, across 500 m of depth. The magnitude of POC flux was not consistently associated with a dominant particle class, but particle classes did influence flux attenuation. Long fecal pellets attenuated most rapidly with depth whereas certain other classes attenuated little or not at all with depth. Small particles (<100 µm) consistently contributed ∼5% to total POC flux in samples with higher magnitude fluxes. The relative importance of these small particle classes (spherical mini pellets, short oval fecal pellets, and dense detritus) increased in low flux environments (up to 46% of total POC flux). Imaging approaches that resolve large variations in particle composition across ocean basins, depth, and time will help to better parameterize biological carbon pump models.

Assessment of holographic microscopy for quantifying marine particle size and concentration.

Holographic microscopy has emerged as a tool for in situ imaging of microscopic organisms and other particles in the marine environment: appealing because of the relatively larger sampling volume and simpler optical configuration compared to other imaging systems. However, its quantitative capabilities have so far remained uncertain, in part because hologram reconstruction and image recognition have required manual operation. Here, we assess the quantitative skill of our automated hologram processing pipeline (CCV Pipeline), to evaluate the size and concentration measurements of environmental and cultured assemblages of marine plankton particles, and microspheres. Over 1 million particles, ranging from 10 to 200 µm in equivalent spherical diameter, imaged by the 4-Deep HoloSea digital inline holographic microscope (DIHM) are analyzed. These measurements were collected in parallel with a FlowCam (FC), Imaging FlowCytobot (IFCB), and manual microscope identification. Once corrections for particle location and nonuniform illumination were developed and applied, the DIHM showed an underestimate in ESD of about 3% to 10%, but successfully reproduced the size spectral slope from environmental samples, and the size distribution of cultures (Dunaliella tertiolecta, Heterosigma akashiwo, and Prorocentrum micans) and microspheres. DIHM concentrations (order 1 to 1000 particles ml-1) showed a linear agreement (r 2 = 0.73) with the other instruments, but individual comparisons at times had large uncertainty. Overall, we found the DIHM and the CCV Pipeline required extensive manual correction, but once corrected, provided concentration and size estimates comparable to the other imaging systems assessed in this study. Holographic cameras are mechanically simple, autonomous, can operate at very high pressures, and provide a larger sampling volume than comparable lens-based tools. Thus, we anticipate that these characterization efforts will be rewarded with novel discovery in new oceanic environments.

Marine plankton community and net primary production responding to island-trapped waves in a stratified oligotrophic ecosystem.

The oligotrophic Adriatic Sea is characterized during a typical summer by low productivity caused by strong water column stratification, which inhibits vertical mixing and nutrient supply to the euphotic zone. These conditions can be disrupted by transient physical forcing, which enhances nutrient fluxes and creates localized hotspots of relatively high net primary production. In this study, plankton abundance and diversity were investigated in relation to the physical forcing and nutrient concentrations in an area affected by island-trapped waves (ITWs) near Lastovo Island (Adriatic Sea). The episodic ITW events resulted in enhanced uplift and vertical excursion of the thermocline, marked by anomalously higher nutrient concentrations and a corresponding increase in net primary production in the thermocline layer. Physicochemical properties explained 11.7 % (p = 0.002) of the variability in micro- and nanophytoplankton and 88.9 % (p = 0.001) in the picoplankton community. A significant response to the ITW phenomenon in the plankton community composition (p = 0.001) was observed for bacterioplankton. Among the identified amplicon sequence variances, primary producers were scarce and mainly represented cyanobacteria (Synechococcus strain CC9902), stramenopiles (Pelagomonas), and chlorophytes (Ostreococcus). The remaining amplicon sequence variances were assigned to the classes Copepoda, parasitic fungi (Meyerozyma spp.), mixotrophic dinoflagellates (family Peridiniales, mostly the genus Blastodinium), and parasitic Ciliophora (Scuticociliata). Bacterial ecological functions corresponded to chemoheterotrophic, degradation, and fermentation processes, whereas samples collected after the most intense ITW episode also showed abundant bacteria linked to microplastic degradation and parasitosis. These results highlight the ecological role of localized physical phenomena in enhancing nearshore primary productivity and fine shifts in plankton taxa in oligotrophic systems.

Tracing the path of carbon export in the ocean though DNA sequencing of individual sinking particles.

Surface phytoplankton communities were linked with the carbon they export into the deep ocean by comparing 18 S rRNA gene sequence communities from surface seawater and individually isolated sinking particles. Particles were collected in sediment traps deployed at locations in the North Pacific subtropical gyre and the California Current. DNA was isolated from individual particles, bulk-collected trap particles, and the surface seawater. The relative sequence abundance of exported phytoplankton taxa in the surface water varied across functional groups and ecosystems. Of the sequences detected in sinking particles, about half were present in large (>300 µm), individually isolated particles and primarily belonged to taxa with small cell sizes (<50 µm). Exported phytoplankton taxa detected only in bulk trap samples, and thus presumably packaged in the smaller sinking size fraction, contained taxa that typically have large cell sizes (>500 µm). The effect of particle degradation on the detectable 18 S rRNA gene community differed across taxa, and differences in community composition among individual particles from the same location largely reflected differences in relative degradation state. Using these data and particle imaging, we present an approach that incorporates genetic diversity into mechanistic models of the ocean's biological carbon pump, which will lead to better quantification of the ocean's carbon cycle.

Sinking flux of particulate organic matter in the oceans: Sensitivity to particle characteristics.

The sinking of organic particles produced in the upper sunlit layers of the ocean forms an important limb of the oceanic biological pump, which impacts the sequestration of carbon and resupply of nutrients in the mesopelagic ocean. Particles raining out from the upper ocean undergo remineralization by bacteria colonized on their surface and interior, leading to an attenuation in the sinking flux of organic matter with depth. Here, we formulate a mechanistic model for the depth-dependent, sinking, particulate mass flux constituted by a range of sinking, remineralizing particles. Like previous studies, we find that the model does not achieve the characteristic 'Martin curve' flux profile with a single type of particle, but instead requires a distribution of particle sizes and/or properties. We consider various functional forms of remineralization appropriate for solid/compact particles, and aggregates with an anoxic or oxic interior. We explore the sensitivity of the shape of the flux vs. depth profile to the choice of remineralization function, relative particle density, particle size distribution, and water column density stratification, and find that neither a power-law nor exponential function provides a definitively superior fit to the modeled profiles. The profiles are also sensitive to the time history of the particle source. Varying surface particle size distribution (via the slope of the particle number spectrum) over 3 days to represent a transient phytoplankton bloom results in transient subsurface maxima or pulses in the sinking mass flux. This work contributes to a growing body of mechanistic export flux models that offer scope to incorporate underlying dynamical and biological processes into global carbon cycle models.

Eddy-driven subduction exports particulate organic carbon from the spring bloom.

The export of particulate organic carbon (POC) from the surface ocean to depth is traditionally ascribed to sinking. Here, we show that a dynamic eddying flow field subducts surface water with high concentrations of nonsinking POC. Autonomous observations made by gliders during the North Atlantic spring bloom reveal anomalous features at depths of 100 to 350 meters with elevated POC, chlorophyll, oxygen, and temperature-salinity characteristics of surface water. High-resolution modeling reveals that during the spring transition, intrusions of POC-rich surface water descend as coherent, 1- to 10-kilometer-scale filamentous features, often along the perimeter of eddies. Such a submesoscale eddy-driven flux of POC is unresolved in global carbon cycle models but can contribute as much as half of the total springtime export of POC from the highly productive subpolar oceans.

Measuring Fluorescent Dye in the Bubbly and Sediment-Laden Surfzone.

Decisions about recreational beach closures would be enhanced if better estimates of surfzone contaminant transport and dilution were available. In situ methods for measuring fluorescent Rhodamine WT dye tracer in the surfzone are presented, increasing the temporal and spatial resolution over previous surfzone techniques. Bubbles and sand suspended by breaking waves in the surfzone interfere with in situ optical fluorometer dye measurements, increasing the lower bound for dye detection ( approximately 1 ppb) and reducing (quenching) measured dye concentrations. Simultaneous turbidity measurements are used to estimate the level of bubble and sand interference and correct dye estimates. After correction, root-mean-square dye concentration errors are estimated to be < 5% of dye concentration magnitude, thus demonstrating the viability of in situ surfzone fluorescent dye measurements. The surfzone techniques developed here may be applicable to other environments with high bubble and sand concentrations (e.g., cascading rivers and streams).