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The marine compound dimethyl sulfoxide (DMSO) is ubiquitous in the world’s surface ocean, constituting one of the largest sources of reduced organic sulfur in seawater. DMSO cycling has been linked to the formation of the climate-active gas dimethyl sulfide (DMS) through a reductive pathway, but the underlying physiological role of DMSO reduction, and the environmental controls on this pathway, remain unresolved. Here we present evidence that DMSO reduction to DMS serves an antioxidant role in phytoplankton through a secondary electron-scavenging pathway that can dissipate excess light-harvested energy, and potentially mitigate the formation of reactive oxygen species (ROS). Results from isotopic tracer experiments demonstrate significant increases in DMSO reduction rates in low-light acclimated natural phytoplankton assemblages exposed to high irradiance. Increased DMSO reduction rates were negatively correlated with non-photochemical quenching, while treatment with the photosynthetic electron transport inhibitor DCMU significantly decreased DMSO reduction, indicating a link to photosynthetically-derived electrons. Our results show that photic stress drives enhanced DMSO reduction in marine phytoplankton, linking DMS production to irradiance and vertical mixing through an electron scavenging mechanism that could serve an antioxidant role.

Iron availability directly affects photosynthesis and limits phytoplankton growth over vast oceanic regions. For this reason, the availability of iron is a crucial variable to consider in the development of active chlorophyll a fluorescence based estimates of phytoplankton primary productivity. These bio-optical approaches require a conversion factor to derive ecologically-relevant rates of CO2-assimilation from estimates of electron transport in photosystem II. The required conversion factor varies significantly across phytoplankton taxa and environmental conditions, but little information is available on its response to iron limitation. In this study, we examine the role of iron limitation, and the interacting effects of iron and light availability, on the coupling of photosynthetic electron transport and CO2-assimilation in marine phytoplankton. Our results show that excess irradiance causes increased decoupling of carbon fixation and electron transport, particularly under iron limiting conditions. We observed that reaction center II specific rates of electron transport (ETR(RCII), mol e- mol RCII(-1) s(-1)) increased under iron limitation, and we propose a simple conceptual model for this observation. We also observed a strong correlation between the derived conversion factor and the expression of non-photochemical quenching. Utilizing a dataset from in situ phytoplankton assemblages across a coastal--oceanic transect in the Northeast subarctic Pacific, this relationship was used to predict ETR(RCII): CO2-assimilation conversion factors and carbon-based primary productivity from FRRF data, without the need for any additional measurements.

The potential interactive effects of iron (Fe) limitation and Ocean Acidification in the Southern Ocean (SO) are largely unknown. Here we present results of a long-term incubation experiment investigating the combined effects of CO2 and Fe availability on natural phytoplankton assemblages from the Weddell Sea, Antarctica. Active Chl a fluorescence measurements revealed that we successfully cultured phytoplankton under both Fe-depleted and Fe-enriched conditions. Fe treatments had significant effects on photosynthetic efficiency (Fv/Fm; 0.3 for Fe-depleted and 0.5 for Fe-enriched conditions), non-photochemical quenching (NPQ), and relative electron transport rates (rETR). pCO2 treatments significantly affected NPQ and rETR, but had no effect on Fv/Fm. Under Fe limitation, increased pCO2 had no influence on C fixation whereas under Fe enrichment, primary production increased with increasing pCO2 levels. These CO2-dependent changes in productivity under Fe-enriched conditions were accompanied by a pronounced taxonomic shift from weakly to heavily silicified diatoms (i.e. from Pseudo-nitzschia sp. to Fragilariopsis sp.). Under Fe-depleted conditions, this functional shift was absent and thinly silicified species dominated all pCO2 treatments (Pseudo-nitzschia sp. and Synedropsis sp. for low and high pCO2, respectively). Our results suggest that Ocean Acidification could increase primary productivity and the abundance of heavily silicified, fast sinking diatoms in Fe-enriched areas, both potentially leading to a stimulation of the biological pump. Over much of the SO, however, Fe limitation could restrict this possible CO2 fertilization effect.

Bottom waters of the northeast Pacific continental shelf naturally experience localized hypoxic conditions, with significant influences on food webs and biogeochemical cycling. In August 2021, extreme hypoxia was detected from several measurement platforms along the southern British Columbia continental shelf, with oxygen concentration <60 μmol kg−1, and a difference from the seasonal climatology of more than 2 standard deviations. Early and intense remote upwelling and local density shifts were associated with an anomalously strong spring phytoplankton bloom, which likely stimulated localized respiration of subsurface organic matter. This event was concurrent with unsuitable habitat for Pacific halibut and calcite and aragonite undersaturation throughout most of the water column. The drivers of this extreme low oxygen event could be enhanced under future climate change, with potentially significant impacts on marine ecology and biogeochemistry. Plain Language Summary Most marine organisms consume oxygen, and are therefore impacted when seawater oxygen concentrations reach low values. Extreme low oxygen concentrations are rare in the coastal waters of British Columbia, Canada. However, unusually strong oxygen depletion was observed off the coast of Vancouver Island during summer 2021. Unusually strong, early season upwelling winds along the California coast impacted Vancouver Island by causing nutrient‐rich water to be mixed into the surface, stimulating a large and earlier than usual spring phytoplankton bloom. Decomposition of the bloom‐derived organic carbon consumed local subsurface oxygen throughout the spring and summer. These subtle changes in timing and intensity of seasonal processes likely caused this low oxygen event, which was also associated with high concentrations of inorganic carbon, leading to ocean acidification. Such extreme low oxygen events, even if short‐lived, can have a significant impact on marine ecosystems, restricting the habitat available for groundfish species, such as Pacific halibut, and impacting the formation of carbonate shells by various organisms. The drivers of extreme low oxygen events are projected to intensify as climate change progresses. Key Points Widespread extreme hypoxia was observed throughout the water column along the southern British Columbia continental shelf in summer 2021 Early and intense upwelling followed by a strong biological response contributed to oxygen depletion in subsurface waters Multiple indices suggest that this extreme event was concurrent with unsuitable habitat conditions for groundfish and calcifying organisms

We report the first measurements of coupled nitrogen (N) and oxygen (O) isotope fractionation of nitrate by laboratory cultures of denitrifying bacteria. Two seawater strains (Pseudomonas stutzeri, Ochrobactrum sp.) and three freshwater strains (Paracoccus denitrificans, Pseudomonas chlororaphis, Rhodobacter sphaeroides) were examined. Among four strains of facultative anaerobic denitrifiers, N and O isotope effects were variable, ranging from 5‰ to 25‰, with evidence for a drop in the isotope effects as nitrate concentrations approached the half- saturation constant for nitrate transport. O isotope effects were similar to their corresponding N isotope effect, such that the progressive increase in nitrate δ¹⁸O, when plotted against that in¹⁵N (where${\\rm{\\delta }}{}^{18}{\\rm{O}}_{{\\rm{sample}}} = [({}^{18}{\\rm{O:}}\\,{}^{16}{\\rm{O)}}_{{\\rm{sample}}} /({}^{18}{\\rm{O:}}\\,{}^{16}{\\rm{O)}}_{{\\rm{reference}}} - 1] \\times \\,1000$, and ${\\rm{\\delta }}^{15} {\\rm{N}}_{{\\rm{sample}}} = [({}^{15}{\\rm{N:}}\\,{}^{14}{\\rm{N)}}_{{\\rm{sample}}} /({}^{15}{\\rm{N:}}\\,{}^{14}{\\rm{N}})_{{\\rm{reference}}} - 1] \\times 1000)$, yielded slopes of 0.86 to 1.02, with a mean value of 0.96. R. sphaeroides, a photo-heterotroph that possesses only a periplasmic (nonrespiring) dissimilatory nitrate reductase, showed less variability in nitrate N isotope effects, between 13‰ and 20‰, with a modal value of ~15‰. In contrast to the respiratory denitrifiers, R. sphaeroides consistently showed a distinct ratio of δ¹⁸O to δ¹⁵N change of 0~.62. We hypothesize that heavy N and O isotope discrimination during respiratory denitrification occurs during the intracellular reduction of nitrate by the respiratory nitrate reductase, and the observed magnitude of fractionation is likely regulated by the ratio of cellular nitrate efflux relative to uptake. The data for R. sphaeroides are consistent with isotope discrimination directly reflecting the N and O isotope effects of the periplasmic nitrate reductase NAP, without modification by nitrate uptake and efflux.

We employed Fast Repetition Rate fluorometry for high-resolution mapping of marine phytoplankton photophysiology and primary photochemistry in the Lancaster Sound and Barrow Strait regions of the Canadian Arctic Archipelago in the summer of 2019. Continuous ship-board analysis of chlorophyll a variable fluorescence demonstrated relatively low photochemical efficiency over most of the cruise-track, with the exception of localized regions within Barrow Strait, where there was increased vertical mixing and proximity to land-based nutrient sources. Along the full transect, we observed strong non-photochemical quenching of chlorophyll fluorescence, with relaxation times longer than the 5-minute period used for dark acclimation. Such long-term quenching effects complicate continuous underway acquisition of fluorescence amplitude-based estimates of photosynthetic electron transport rates, which rely on dark acclimation of samples. As an alternative, we employed a new algorithm to derive electron transport rates based on analysis of fluorescence relaxation kinetics, which does not require dark acclimation. Direct comparison of kinetics- and amplitude-based electron transport rate measurements demonstrated that kinetic-based estimates were, on average, 2-fold higher than amplitude-based values. The magnitude of decoupling between the two electron transport rate estimates increased in association with photophysiological diagnostics of nutrient stress. Discrepancies between electron transport rate estimates likely resulted from the use of different photophysiological parameters to derive the kinetics- and amplitude-based algorithms, and choice of numerical model used to fit variable fluorescence curves and analyze fluorescence kinetics under actinic light. Our results highlight environmental and methodological influences on fluorescence-based photochemistry estimates, and prompt discussion of best-practices for future underway fluorescence-based efforts to monitor phytoplankton photosynthesis.

April 22, 2020, marks the 50th anniversary of Earth Day and the birth of the modern environmental movement. As we look back over the past half century, we can gain significant insights into the evolving human imprint on Earth’s biophysical systems, and the role of science and scientists in driving societal transitions toward greater sustainability. Science is a foundation for such transitions, but it is not enough. Rather, it is through wide collaborations across fields, including law, economics, and politics, and through direct engagement with civil society, that science can illuminate a better path forward. This is illustrated through a number of case studies highlighting the role of scientists in leading positive societal change, often in the face of strong oppositional forces. The past five decades reveal significant triumphs of environmental protection, but also notable failures, which have led to the continuing deterioration of Earth’s natural systems. Today, more than ever, these historical lessons loom large as we face increasingly complex and pernicious environmental problems.

The Arctic and subarctic shelf seas, which sustain large fisheries and contribute to global biogeochemical cycling, are particularly sensitive to ongoing ocean acidification (that is, decreasing seawater pH due to anthropogenic CO2 emissions). Yet, little information is available on the effects of ocean acidification on natural phytoplankton assemblages, which are the main primary producers in high-latitude waters. Here we show that coastal Arctic and subarctic primary production is largely insensitive to ocean acidification over a large range of light and temperature levels in different experimental designs. Out of ten CO2-manipulation treatments, significant ocean acidification effects on primary productivity were observed only once (at temperatures below 2 °C), and shifts in the species composition occurred only three times (without correlation to specific experimental conditions). These results imply a high capacity to compensate for environmental variability, which can be understood in light of the environmental history, tolerance ranges and intraspecific diversity of the dominant phytoplankton species.

Active chlorophyll a fluorescence approaches, including fast repetition rate fluorometry (FRRF), have the potential to provide estimates of phytoplankton primary productivity at an unprecedented spatial and temporal resolution. FRRF-derived productivity rates are based on estimates of charge separation in reaction center II (ETRRCII), which must be converted into ecologically relevant units of carbon fixation. Understanding sources of variability in the coupling of ETRRCII and carbon fixation provides physiological insight into phytoplankton photosynthesis and is critical for the application of FRRF as a primary productivity measurement tool. In the present study, we simultaneously measured phytoplankton carbon fixation and ETRRCII in the iron-limited NE subarctic Pacific over the course of a diurnal cycle. We show that rates of ETRRCII are closely tied to the diurnal cycle in light availability, whereas rates of carbon fixation appear to be influenced by endogenous changes in metabolic energy allocation under iron-limited conditions. Unsynchronized diurnal oscillations of the two rates led to 3.5-fold changes in the conversion factor between ETRRCII and carbon fixation (Kc / nPSII). Consequently, diurnal variability in phytoplankton carbon fixation cannot be adequately captured with FRRF approaches if a constant conversion factor is applied. Utilizing several auxiliary photophysiological measurements, we observed that a high conversion factor is associated with conditions of excess light and correlates with the increased expression of non-photochemical quenching (NPQ) in the pigment antenna, as derived from FRRF measurements. The observed correlation between NPQ and Kc / nPSII requires further validation but has the potential to improve estimates of phytoplankton carbon fixation rates from FRRF measurements alone.

High‐latitude oceans are areas of high primary production despite temperatures that are often well below the thermal optima of enzymes, including the key Calvin Cycle enzyme, Ribulose 1,5 bisphosphate carboxylase oxygenase (Rubisco). We measured carbon fixation rates, protein content and Rubisco abundance and catalytic rates during an intense diatom bloom in the Western Antarctic Peninsula (WAP) and in laboratory cultures of a psychrophilic diatom (Fragilariopsis cylindrus). At −1°C, the Rubisco turnover rate, kcₐₜᶜ, was 0.4 C s⁻¹per site and the half saturation constant for CO₂was 15 μM (vs c. 3 C s⁻¹per site and 50 μM at 20°C). To achieve high carboxylation rates, psychrophilic diatoms increased Rubisco abundance to c. 8% of biomass (vs c. 0.6% at 20°C), along with their total protein content, resulting in a low carbon : nitrogen ratio of c. 5. In psychrophilic diatoms, Rubisco must be almost fully active and near CO₂saturation to achieve carbon fixation rates observed in the WAP. Correspondingly, total protein concentrations were close to the highest ever measured in phytoplankton and likely near the maximum possible. We hypothesize that this high protein concentration, like that of Rubisco, is necessitated by slow enzyme rates, and that carbon fixation rates in the WAP are near a theoretical maximum.