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Southern Ocean primary productivity is principally controlled by adjustments in light and iron limitation, but the spatial and temporal determinants of iron availability, accessibility, and demand are poorly constrained, which hinders accurate long-term projections. We present a multidecadal record of phytoplankton photophysiology between 1996 and 2022 from historical in situ datasets collected by Biogeochemical Argo (BGC-Argo) floats and ship-based platforms. We find a significant multidecadal trend in irradiance-normalized nonphotochemical quenching due to increasing iron stress, with concomitant declines in regional net primary production. The observed trend of increasing iron stress results from changing Southern Ocean mixed-layer physics as well as complex biological and chemical feedback that is indicative of important ongoing changes to the Southern Ocean carbon cycle.

We studied phytoplankton and protozooplankton community composition based on light microscopy, flow cytometry, and photosynthetic pigment data in the Atlantic sector of the Southern Ocean during March 2019 (early austral autumn). Sampling was focused on the area east of the prime meridian in the Kong Håkon VII Hav, including Astrid Ridge, Maud Rise, and a south–north transect at 6∘ E. Phytoplankton community composition throughout the studied area was characterized by oceanic diatoms typical of the iron-depleted high-nutrient, low-chlorophyll (HNLC) Southern Ocean. Topography and wind-driven iron supply likely sustained blooms dominated by the centric diatom Chaetoceros dichaeta at Maud Rise and at a station north of the 6∘ E transect. For the remainder of the 6∘ E transect, diatom composition was similar to the previously mentioned bloom stations, but flagellates dominated in abundance, suggesting a post-bloom situation and likely top-down control by krill on the bloom-forming diatoms. Among flagellates, species with haptophyte-type pigments were the dominating group. At Astrid Ridge, overall abundances were lower and pennate diatoms were more numerous than centric diatoms, but the community composition was nevertheless typical of HNLC areas. The observations described here show that C. dichaeta can form blooms beyond the background biomass level and also fuels both carbon export and upper trophic levels within HNLC areas. This study is the first thorough assessment of phytoplankton communities in this region and can be compared to other seasons in future studies.

Quantifying the strength and efficiency of the Southern Ocean biological carbon pump (BCP) and its response to predicted changes in the Earth's climate is fundamental to our ability to predict long-term changes in the global carbon cycle and, by extension, the impact of continued anthropogenic perturbation of atmospheric CO 2 . There is little agreement, however, in climate model projections of the sensitivity of the Southern Ocean BCP to climate change, with a lack of consensus in even the direction of predicted change, highlighting a gap in our understanding of a major planetary carbon flux. In this review, we summarize relevant research that highlights the important role of fine-scale dynamics (both temporal and spatial) that link physical forcing mechanisms to biogeochemical responses that impact the characteristics of the seasonal cycle of phytoplankton and by extension the BCP. This approach highlights the potential for integrating autonomous and remote sensing observations of fine scale dynamics to derive regionally optimized biogeochemical parameterizations for Southern Ocean models. Ongoing development in both the observational and modelling fields will generate new insights into Southern Ocean ecosystem function for improved predictions of the sensitivity of the Southern Ocean BCP to climate change. This article is part of a discussion meeting issue 'Heat and carbon uptake in the Southern Ocean: the state of the art and future priorities'.

The High Nutrient-Low Chlorophyll condition of the Southern Ocean is generally thought to be caused by the low bioavailability of micronutrients, particularly iron, which plays an integral role in phytoplankton photosynthesis. Nevertheless, the Southern Ocean experiences seasonal blooms that generally initiate in austral spring, peak in summer and extend into autumn. This seasonal increase in primary productivity is typically linked to the seasonal characteristics of nutrient and light supply. To better understand the constraints on productivity in the Antarctic Sea-Ice Zone (SIZ), the photophysiological response of phytoplankton to iron addition was investigated during autumn along the Antarctic coast off Dronning Maud Land. Five short-term (24 hr) incubation experiments were conducted around Astrid Ridge (68° S) and along a 6° E transect, where an autumn bloom was identified in the region of the western SIZ. Surface iron concentrations ranged from 0.27 to 1.39 nM around Astrid Ridge, and 0.56 to 0.63 nM along the 6° E transect. The photophysiological response of phytoplankton to iron addition, measured through the photosynthetic efficiency and the absorption cross-section for photosystem II, showed no significant responses. This confirms that phytoplankton were not iron-limited at the time and that ambient iron concentrations were sufficient to fulfil the cellular requirements. This provides new insights into extended iron replete post-bloom conditions in the typically assumed iron deficient High Nutrient-Low Chlorophyll Southern Ocean.

The uptake and application of single turnover chlorophyll fluorometers to the study of phytoplankton ecosystem status and microbial functions has grown considerably in the last two decades. However, standardization of measurement protocols, processing of fluorescence transients and quality control of derived photosynthetic parameters is still lacking and makes community goals of large global databases of high-quality data unrealistic. We introduce the Python package Phytoplankton Photophysiology Utilities (PPU), an adaptable and open-source interface between Fast Repetition Rate and Fluorescence Induction and Relaxation instruments and python. The PPU package includes a variety of functions for the loading, processing and quality control of single turnover fluorescence transients from many commercially available instruments. PPU provides the user with greater flexibility in the application of the Kolber-Prasil-Falkowski model; tools for plotting, quality control, correcting instrument biases and high-throughput processing with ease; and a greater appreciation for the uncertainties in derived photosynthetic parameters. Using data from three research cruises across different biogeochemical regimes, we provide example applications of PPU to fit raw active chlorophyll-a fluorescence data from three commercial instruments and demonstrate tools which help to reduce uncertainties in the final fitted parameters.

Knowing the magnitude and timing of pelagic primary production is important for ecosystem and carbon sequestration studies, in addition to providing basic understanding of phytoplankton functioning. In this study we use data from an ecosystem cruise to Kong Håkon VII Hav, in the Atlantic sector of the Southern Ocean, in March 2019 and more than two decades of satellite-derived ocean color to study phytoplankton bloom phenology. During the cruise we observed phytoplankton blooms in different bloom phases. By correlating bloom phenology indices (i.e., bloom initiation and end) based on satellite remote sensing to the timing of changes in environmental conditions (i.e., sea ice, light, and mixed layer depth) we studied the environmental factors that seemingly drive phytoplankton blooms in the area. Our results show that blooms mainly take place in January and February, consistent with previous studies that include the area. Sea ice retreat controls the bloom initiation in particular along the coast and the western part of the study area, whereas bloom end is not primarily connected to sea ice advance. Light availability in general is not appearing to control the bloom termination, neither is nutrient availability based on the autumn cruise where we observed non-depleted macronutrient reservoirs in the surface. Instead, we surmise that zooplankton grazing plays a potentially large role to end the bloom, and thus controls its duration. The spatial correlation of the highest bloom magnitude with marked topographic features indicate that the interaction of ocean currents with sea floor topography enhances primary productivity in this area, probably by natural fertilization. Based on the bloom timing and magnitude patterns, we identified five different bloom regimes in the area. A more detailed understanding of the region will help to highlight areas with the highest relevance for the carbon cycle, the marine ecosystem and spatial management. With this gained understanding of bloom phenology, it will also be possible to study potential shifts in bloom timing and associated trophic mismatch caused by environmental changes.

The seasonal and sub-seasonal dynamics of iron availability within the sub-Antarctic zone (SAZ; ∼40–45∘ S) play an important role in the distribution, biomass and productivity of the phytoplankton community. The variability in iron availability is due to an interplay between winter entrainment, diapycnal diffusion, storm-driven entrainment, atmospheric deposition, iron scavenging and iron recycling processes. Biological observations utilizing grow-out iron addition incubation experiments were performed at different stages of the seasonal cycle within the SAZ to determine whether iron availability at the time of sampling was sufficient to meet biological demands at different times of the growing season. Here we demonstrate that at the beginning of the growing season, there is sufficient iron to meet the demands of the phytoplankton community, but that as the growing season develops the mean iron concentrations in the mixed layer decrease and are insufficient to meet biological demand. Phytoplankton increase their photosynthetic efficiency and net growth rates following iron addition from midsummer to late summer, with no differences determined during early summer, suggestive of seasonal iron depletion and an insufficient resupply of iron to meet biological demand. The result of this is residual macronutrients at the end of the growing season and the prevalence of the high-nutrient low-chlorophyll (HNLC) condition. We conclude that despite the prolonged growing season characteristic of the SAZ, which can extend into late summer/early autumn, results nonetheless suggest that iron supply mechanisms are insufficient to maintain potential maximal growth and productivity throughout the season.

Copyright: 2017 Wiley. Due to copyright restrictions, the attached PDF file only contains the abstract of the full text item. For access to the full text item, kindly consult the publisher's website.

Studying the biogeochemistry of the Southern Ocean using remote sensing relies on accurate interpretation of ocean colour through bio-optical and biogeochemical relationships between quantities and properties of interest. During the Antarctic Circumnavigation Expedition of the 2016/2017 Austral Summer, we collected a spatially comprehensive dataset of phytoplankton pigment concentrations, particulate absorption and particle size distribution and compared simple bio-optical and particle property relationships as a function of chlorophyll a. Similar to previous studies we find that the chlorophyll-specific phytoplankton absorption coefficient is significantly lower than in other oceans at comparable chlorophyll concentrations. This appears to be driven in part by lower concentrations of accessory pigments per unit chlorophyll a as well as increased pigment packaging due to relatively larger sized phytoplankton at low chlorophyll a than is typically observed in other oceans. We find that the contribution of microphytoplankton (>20 µm size) to chlorophyll a estimates of phytoplankton biomass is significantly higher than expected for the given chlorophyll a concentration, especially in higher latitudes south of the Southern Antarctic Circumpolar Current Front. Phytoplankton pigments are more packaged in larger cells, which resulted in a flattening of phytoplankton spectra as measured in these samples when compared to other ocean regions with similar chlorophyll a concentration. Additionally, we find that at high latitude locations in the Southern Ocean, pheopigment concentrations can exceed mono-vinyl chlorophyll a concentrations. Finally, we observed very different relationships between particle volume and chlorophyll a concentrations in high and low latitude Southern Ocean waters, driven by differences in phytoplankton community composition and acclimation to environmental conditions and varying contribution of non-algal particles to the particulate matter. Our data confirm that, as previously suggested, the relationships between bio-optical properties and chlorophyll a in the Southern Ocean are different to other oceans. In addition, distinct bio-optical properties were evident between high and low latitude regions of the Southern Ocean basin. Here we provide a region-specific set of power law functions describing the phytoplankton absorption spectrum as a function of chlorophyll a.

Seasonal progression of dissolved iron (DFe) concentrations in the upper water column were examined during four occupations in the Atlantic sector of the Southern Ocean. DFe inventories from euphotic and aphotic reservoirs decreased progressively from July to February, while dissolved inorganic nitrogen (DIN) decreased from July to January with no significant change between January and February. Results suggest that between July and January, DFe loss from both euphotic and aphotic reservoirs were predominantly in support of phytoplankton growth (Iron to carbon (Fe:C) uptake ratio of 16±3 μmol mol‐1) highlighting the importance of the “winter DFe‐reservoir” for biological uptake. During January to February, excess loss of DFe relative to DIN (Fe:C uptake ratio of 44±8 μmol mol‐1 and aphotic DFe loss rate of 0.34±0.06 μmol m‐2 d‐1) suggests that scavenging is the dominant removal mechanism of DFe from the aphotic, while continued production is likely supported by recycled nutrients. Plain Language Summary Trace metal iron is one of the limiting nutrients for primary productivity in the Southern Ocean; however the relative importance of seasonal iron supply and sinks remains poorly understood, due to sparse data coverage across the seasonal cycle and lack of high‐resolution dissolved iron (DFe) measurements. Here, we present four “snap‐shots” of DFe measurements at a single station in the south‐east Southern Atlantic Ocean (one in winter and three in late spring‐summer), to address the seasonal evolution of DFe and dissolved inorganic nitrogen (DIN) concentrations within the biologically active sunlit and subsurface reservoirs. We observed a seasonal depletion of DFe inventories from July‐February, while DIN inventories decreases from July‐January with no concomitant changes between January‐February. This suggests that, in addition to biological uptake in the sunlit layer, the observed decrease in DFe inventories below this (relative to DIN) is driven by aggregation and incorporation of iron particles into larger "marine snow" sinking particles, while nutrient recycling is driving the observed continuation of primary productivity during late summer. Our results provide insight into seasonal change of DFe availability in different reservoirs where interplay between removal and supply processes are controlling its distributions and bioavailability to support upper surface primary production.

The Ross Sea is one of the most productive regions in the Southern Ocean, with a significant role in carbon cycling as well as the massive abundance of higher trophic levels. The seasonal cycle is well established with an early summer Phaeocystis antarctica bloom that declines followed by a diatom bloom in late summer. This seasonal progression of the phytoplankton has been linked to the availability of iron in the mixed layer. Investigating the temporal progression of iron limitation is often limited by both the decreased sampling resolution from traditional platforms, such as ships, and the lack of regular deployments of specific sensors that can measure phytoplankton physiology. Through the use of a novel technique that uses the degree of quenching (NPQGlider), determined from a standard fluorometer deployed on a buoyancy glider, a proxy for iron limitation, αNPQ, was calculated for a glider time series in the Ross Sea from December 2011 to February 2012. Surface chlorophyll concentrations indicated that there were four stages: the first being a pre-bloom phase, the second in which phytoplankton growth was rapid, resulting in the accumulation of biomass; the third in which biomass in the surface layer decreased, and the fourth in which chlorophyll concentrations remained low but the POC:Chl ratio increased. The levels of NPQGlider in this region were much higher compared to other Southern Ocean regions, with the highest levels in the third phase. Similarly, αNPQ remained low throughout most of the time series except for the transition from the second to third phase when the surface biomass decreases. The increase in POC:Chl ratios in the final phase combined with the low values of αNPQ suggest the switch from a Phaeocystis antarctica bloom to a potentially non-iron limited diatom bloom. These results confirm that the application of novel methodologies to proven and reliable sensors will provide a greater understanding of biogeochemical cycles and their controls throughout the ocean.

The bioavailability of iron influences the distribution, biomass and productivity of phytoplankton in the Ross Sea, one of the most productive regions in the Southern Ocean. We mapped the spatial and temporal extent and severity of iron-limitation of the native phytoplankton assemblage using long- (> 24 h) and short-term (24 h) iron-addition experiments along with physiological and molecular characterisations during a cruise to the Ross Sea in December–February 2012. Phytoplankton increased their photosynthetic efficiency in response to iron addition, suggesting proximal iron limitation throughout most of the Ross Sea during summer. Molecular and physiological data further indicate that as nitrate is removed from the surface ocean the phytoplankton community transitions to one displaying an iron-efficient photosynthetic strategy characterised by an increase in the size of photosystem II (PSII) photochemical cross section (σ PSII ) and a decrease in the chlorophyll-normalised PSII abundance. These results suggest that phytoplankton with the ability to reduce their photosynthetic iron requirements are selected as the growing season progresses, which may drive the well-documented progression from Phaeocystis antarctica- assemblages to diatom-dominated phytoplankton. Such a shift in the assemblage-level photosynthetic strategy potentially mediates further drawdown of nitrate following the development of iron deficient conditions in the Ross Sea.

Iron availability influences phytoplankton physiology and growth over more than one-third of the surface oceans, with recent evidence even indicating iron stress during and following the latter stages of the spring bloom in the high latitude North Atlantic. The mechanistic basis of the phytoplankton physiological responses used for diagnosing iron stress and the broader ecophysiological consequences of iron stress within natural phytoplankton communities still remain unclear. We describe photosynthetic macromolecular and physiological responses of natural phytoplankton communities both in situ and within factorial nutrient-addition (iron and nitrogen) experiments over a seasonal growth cycle in the subpolar North Atlantic. The abundance of quantified photosynthetic proteins increased under relief of iron stress, with the synthesis of the associated protein catalytic complexes accounting for , 3% of inorganic nitrogen drawdown. However, no significant differences in the stoichiometries of the photosynthetic complexes were observed, suggesting that re-modeling of the photosynthetic electron transport chain was not a significant influence on the community-level ecophysiological responses to iron stress. In marked contrast, iron stress resulted in significant increases in the cellular ratios of chlorophyll to the photosynthetic catalysts, including photosystem II (PSII), alongside a marked increase in PSII normalized chlorophyll fluorescence. Characteristic depressions in apparent photosynthetic energy conversion efficiencies in iron-limited oceanic regions are thus likely driven by a significant accumulation of partially energetically uncoupled chlorophyll-binding complexes. Such iron-stress–induced chlorophyll-binding proteins may contribute , 40% of the total chlorophyll pool during iron-stressed periods.

The Southern Ocean (SO) is highly sensitive to climate change. Therefore, an accurate estimate of phytoplankton biomass is key to being able to predict the climate trajectory of the 21st century. In this study, MODerate resolution Imaging Spectroradiometer (MODIS), on board EOS Aqua spacecraft, Level 2 (nominal 1 km × 1 km resolution) chlorophyll-a (C S a t ) and Particulate Organic Carbon (POC s a t ) products are evaluated by comparison with an in situ dataset from 11 research cruises (2008–2017) to the SO, across multiple seasons, which includes measurements of POC and chlorophyll-a (C i n s i t u ) from both High Performance Liquid Chromatography (C H P L C ) and fluorometry (C F l u o ). Contrary to a number of previous studies, results highlighted good performance of the algorithm in the SO when comparing estimations with HPLC measurements. Using a time window of ±12 h and a mean satellite chlorophyll from a 5 × 5 pixel box centered on the in situ location, the median C S a t :C i n s i t u ratios were 0.89 (N = 46) and 0.49 (N = 73) for C H P L C and C F l u o respectively. Differences between C H P L C and C F l u o were associated with the presence of diatoms containing chlorophyll-c pigments, which induced an overestimation of chlorophyll-a when measured fluorometrically due to a potential overlap of the chlorophyll-a and chlorophyll-c emission spectra. An underestimation of ∼0.13 mg m − 3 was observed for the global POC algorithm. This error was likely due to an overestimate of in situ POC i n s i t u measurements from the impact of dissolved organic carbon not accounted for in the blank correction. These results highlight the important implications of different in situ methodologies when validating ocean colour products.

Aerosol deposition from the 2010 eruption of the Icelandic volcano Eyjafjallajokull resulted in significant dissolved iron (DFe) inputs to the Iceland Basin of the North Atlantic. Unique ship-board measurements indicated strongly enhanced DFe concentrations (up to 10 nM) immediately under the ash plume. Bioassay experiments performed with ash collected at sea under the plume also demonstrated the potential for associated Fe release to stimulate phytoplankton growth and nutrient drawdown. Combining Fe dissolution measurements with modeled ash deposition suggested that the eruption had the potential to increase DFe by >0.2 nM over an area of up to 570,000 km2. Although satellite ocean color data only indicated minor increases in phytoplankton abundance over a relatively constrained area, comparison of in situ nitrate concentrations with historical records suggested that ash deposition may have resulted in enhanced major nutrient drawdown. Our observations thus suggest that the 2010 Eyjafjallajokull eruption resulted in a significant perturbation to the biogeochemistry of the Iceland Basin.

The high-latitude North Atlantic (HLNA) is characterized by a marked seasonal phytoplankton bloom, which removes the majority of surface macronutrients. However, incomplete nitrate depletion is frequently observed during summer in the region, potentially reflecting the seasonal development of an iron (Fe) limited phytoplankton community. In order to investigate the seasonal development and spatial extent of iron stress in the HLNA, nutrient addition experiments were performed during the spring (May) and late summer (July and August) of 2010. Grow-out experiments (48–120 h) confirmed the potential for iron limitation in the region. Short-term (24 h) incubations further enabled high spatial coverage and mapping of phytoplankton physiological responses to iron addition. The difference in the apparent maximal photochemical yield of photosystem II (PSII) (Fv : Fm) between nutrient (iron) amended and control treatments (D(Fv : Fm)) was used as a measure of the relative degree of iron stress. The combined observations indicated variability in the seasonal cycle of iron stress between different regions of the Irminger and Iceland Basins of the HLNA, related to the timing of the annual bloom cycle in contrasting biogeochemical provinces. Phytoplankton iron stress developed during the transition from the prebloom to peak bloom conditions in the HLNA and was more severe for larger cells. Subsequently, iron stress was reduced in regions where macronutrients were depleted following the bloom. Iron availability plays a significant role in the biogeochemistry of the HLNA, potentially lowering the efficiency of one of the strongest biological carbon pumps in the ocean.

Iron availability limits primary production in30% of the world's oceans; hence phytoplankton have developed acclimation strategies. In particular, cyanobacteria express IsiA (iron-stress-induced) under iron stress, which can become the most abundant chl-binding protein in the cell. Within iron-limited oceanic regions with significant cyanobacterial biomass, IsiA may represent a significant fraction of the total chl. We spectroscopically measured the effective cross-section of the photosynthetic reaction center PSI (σPSI ) in vivo and biochemically quantified the absolute abundance of PSI, PSII, and IsiA in the model cyanobacterium Synechocystis sp. PCC 6803. We demonstrate that accumulation of IsiA results in a ∼60% increase in σPSI , in agreement with the theoretical increase in cross-section based on the structure of the biochemically isolated IsiA-PSI supercomplex from cyanobacteria. Deriving a chl budget, we suggest that IsiA plays a primary role as a light-harvesting antenna for PSI. On progressive iron-stress in culture, IsiA continues to accumulate without a concomitant increase in σPSI , suggesting that there may be a secondary role for IsiA. In natural populations, the potential physiological significance of the uncoupled pool of IsiA remains to be established. However, the functional role as a PSI antenna suggests that a large fraction of IsiA-bound chl is directly involved in photosynthetic electron transport.

Iron availability limits primary production in >30% of the world's oceans; hence phytoplankton have developed acclimation strategies. In particular, cyanobacteria express IsiA (iron-stress-induced) under iron stress, which can become the most abundant chl-binding protein in the cell. Within iron-limited oceanic regions with significant cyanobacterial biomass, IsiA may represent a significant fraction of the total chl. We spectroscopically measured the effective cross-section of the photosynthetic reaction center PSI (σPSI ) in vivo and biochemically quantified the absolute abundance of PSI, PSII, and IsiA in the model cyanobacterium Synechocystis sp. PCC 6803. We demonstrate that accumulation of IsiA results in a ∼60% increase in σPSI , in agreement with the theoretical increase in cross-section based on the structure of the biochemically isolated IsiA-PSI supercomplex from cyanobacteria. Deriving a chl budget, we suggest that IsiA plays a primary role as a light-harvesting antenna for PSI. On progressive iron-stress in culture, IsiA continues to accumulate without a concomitant increase in σPSI , suggesting that there may be a secondary role for IsiA. In natural populations, the potential physiological significance of the uncoupled pool of IsiA remains to be established. However, the functional role as a PSI antenna suggests that a large fraction of IsiA-bound chl is directly involved in photosynthetic electron transport.