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1. Seasonal mixed layer depth shapes phytoplankton physiology, viral production, and accumulation in the North Atlantic

   https://doi.org/https://doi.org/10.1038/s41467-021-26836-1  

   Diaz, Ben P.; Knowles, Ben; Johns, Christopher T.; Laber, Christien P.; Bondoc, Karen.G.V.; Haramaty, Liti; Natale, Frank; Harvey, Elizabeth L; Kramer, Sasha J.; Bolaños, Luis M; et al (November 2021, Nature communications)

   Seasonal shifts in phytoplankton accumulation and loss largely follow changes in mixed layer depth, but the impact of mixed layer depth on cell physiology remains unexplored. Here, we investigate the physiological state of phytoplankton populations associated with distinct bloom phases and mixing regimes in the North Atlantic. Stratification and deep mixing alter community physiology and viral production, effectively shaping accumulation rates. Communities in relatively deep, early-spring mixed layers are characterized by low levels of stress and high accumulation rates, while those in the recently shallowed mixed layers in late-spring have high levels of oxidative stress. Prolonged stratification into early autumn manifests in negative accumulation rates, along with pronounced signatures of compromised membranes, death-related protease activity, virus production, nutrient drawdown, and lipid markers indicative of nutrient stress. Positive accumulation renews during mixed layer deepening with transition into winter, concomitant with enhanced nutrient supply and lessened viral pressure. 

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2. Marine phytoplankton downregulate core photosynthesis and carbon storage genes upon rapid mixed layer shallowing

   https://doi.org/10.1038/s41396-023-01416-x  

   Diaz, Ben P.; Zelzion, Ehud; Halsey, Kimberly; Gaube, Peter; Behrenfeld, Michael; Bidle, Kay D. (July 2023, The ISME Journal)

   Abstract Marine phytoplankton are a diverse group of photoautotrophic organisms and key mediators in the global carbon cycle. Phytoplankton physiology and biomass accumulation are closely tied to mixed layer depth, but the intracellular metabolic pathways activated in response to changes in mixed layer depth remain less explored. Here, metatranscriptomics was used to characterize the phytoplankton community response to a mixed layer shallowing (from 233 to 5 m) over the course of two days during the late spring in the Northwest Atlantic. Most phytoplankton genera downregulated core photosynthesis, carbon storage, and carbon fixation genes as the system transitioned from a deep to a shallow mixed layer and shifted towards catabolism of stored carbon supportive of rapid cell growth. In contrast, phytoplankton genera exhibited divergent transcriptional patterns for photosystem light harvesting complex genes during this transition. Active virus infection, taken as the ratio of virus to host transcripts, increased in the Bacillariophyta (diatom) phylum and decreased in the Chlorophyta (green algae) phylum upon mixed layer shallowing. A conceptual model is proposed to provide ecophysiological context for our findings, in which integrated light limitation and lower division rates during transient deep mixing are hypothesized to disrupt resource-driven, oscillating transcript levels related to photosynthesis, carbon fixation, and carbon storage. Our findings highlight shared and unique transcriptional response strategies within phytoplankton communities acclimating to the dynamic light environment associated with transient deep mixing and shallowing events during the annual North Atlantic bloom. 

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3. Under‐Ice Phytoplankton Blooms Inhibited by Spring Convective Mixing in Refreezing Leads

   https://doi.org/10.1002/2016JC012575  

   Lowry, Kate E.; Pickart, Robert S.; Selz, Virginia; Mills, Matthew M.; Pacini, Astrid; Lewis, Kate M.; Joy‐Warren, Hannah L.; Nobre, Carolina; van Dijken, Gert L.; Grondin, Pierre‐Luc; et al (January 2018, Journal of Geophysical Research: Oceans)

   Abstract Spring phytoplankton growth in polar marine ecosystems is limited by light availability beneath ice‐covered waters, particularly early in the season prior to snowmelt and melt pond formation. Leads of open water increase light transmission to the ice‐covered ocean and are sites of air‐sea exchange. We explore the role of leads in controlling phytoplankton bloom dynamics within the sea ice zone of the Arctic Ocean. Data are presented from spring measurements in the Chukchi Sea during the Study of Under‐ice Blooms In the Chukchi Ecosystem (SUBICE) program in May and June 2014. We observed that fully consolidated sea ice supported modest under‐ice blooms, while waters beneath sea ice with leads had significantly lower phytoplankton biomass, despite high nutrient availability. Through an analysis of hydrographic and biological properties, we attribute this counterintuitive finding to springtime convective mixing in refreezing leads of open water. Our results demonstrate that waters beneath loosely consolidated sea ice (84–95% ice concentration) had weak stratification and were frequently mixed below the critical depth (the depth at which depth‐integrated production balances depth‐integrated respiration). These findings are supported by theoretical model calculations of under‐ice light, primary production, and critical depth at varied lead fractions. The model demonstrates that under‐ice blooms can form even beneath snow‐covered sea ice in the absence of mixing but not in more deeply mixed waters beneath sea ice with refreezing leads. Future estimates of primary production should account for these phytoplankton dynamics in ice‐covered waters. 

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4. Synoptic-to-planetary scale wind variability enhances phytoplankton biomass at ocean fronts

   https://doi.org/10.1002/2016JC011899  

   Whitt, D. B.; Taylor, J. R.; Lévy, M. (July 2017, Journal of Geophysical Research: Oceans)

   In nutrient-limited conditions, phytoplankton growth at fronts is enhanced by winds, which drive upward nutrient fluxes via enhanced turbulent mixing and upwelling. Hence, depth-integrated phytoplankton biomass can be 10 times greater at isolated fronts. Using theory and two-dimensional simulations with a coupled physical-biogeochemical ocean model, this paper builds conceptual understanding of the physical processes driving upward nutrient fluxes at fronts forced by unsteady winds with timescales of 4–16 days. The largest vertical nutrient fluxes occur when the surface mixing layer penetrates the nutricline, which fuels phytoplankton in the mixed layer. At a front, mixed layer deepening depends on the magnitude and direction of the wind stress, cross-front variations in buoyancy and velocity at the surface, and potential vorticity at the base of the mixed layer, which itself depends on past wind events. Consequently, mixing layers are deeper and more intermittent in time at fronts than outside fronts. Moreover, mixing can decouple in time from the wind stress, even without other sources of physical variability. Wind-driven upwelling also enhances depth-integrated phytoplankton biomass at fronts; when the mixed layer remains shallower than the nutricline, this results in enhanced subsurface phytoplankton. Oscillatory along-front winds induce both oscillatory and mean upwelling. The mean effect of oscillatory vertical motion is to transiently increase subsurface phytoplankton over days to weeks, whereas slower mean upwelling sustains this increase over weeks to months. Taken together, these results emphasize that wind-driven phytoplankton growth is both spatially and temporally intermittent and depends on a diverse combination of physical processes. 

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5. Net Community Production and Inorganic Carbon Cycling in the Central Irminger Sea

   https://doi.org/10.1029/2024JC021027  

   Yoder, M F; Palevsky, H I; Fogaren, K E (July 2024, Journal of Geophysical Research: Oceans)

   Abstract The subpolar North Atlantic plays an outsized role in the atmosphere‐to‐ocean carbon sink. The central Irminger Sea is home to well‐documented deep winter convection and high phytoplankton production, which drive strong seasonal and interannual variability in regional carbon cycling. We use observational data from moored carbonate chemistry system sensors and annual turn‐around cruise samples at the Ocean Observatories Initiative's Irminger Sea Array to construct a near‐continuous time series of mixed layer total dissolved inorganic carbon (DIC),pCO₂, and total alkalinity from summer 2015 to summer 2022. We use these carbonate chemistry system time series to deconvolve the physical and biological drivers of surface ocean carbon cycling in this region on seasonal, annual, and interannual time scales. We find high annual net community production within the seasonally varying mixed layer, averaging 9.8 ± 1.6 mol m⁻² yr⁻¹with high interannual variability (range of 6.0–13.9 mol m⁻² yr⁻¹). The highest daily net community production rates occur during the late winter and early spring, prior to the observed high chlorophyll concentrations associated with the spring phytoplankton bloom. As a result, the winter and early spring play a much larger role in biological carbon export from the mixed layer than traditionally thought. 

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