Total Water Column Analysis Shows the Importance of a Single Species in Subsurface Chlorophyll Maximum Thin Layers in Stratified Waters

Marine phytoplankton form the base of marine food webs and are the driving force of the marine carbon cycle, so understanding the dynamics of their blooms is critical. While near-surface marine productivity (<10 m water depths) is extensively documented, that of the subsurface is less well characterised. Increasing evidence of the importance of subsurface chlorophyll maxima (SCM) and climatically driven increases in stratification of surface waters that promote SCM development call for improved sampling of the subsurface. To address this, we targeted the summer stratified waters of the Western English Channel, part of the NW European shelf seas, where SCM are commonly developed. In situ holography was applied to undertake the highest ever resolution, total water column, quantitative analysis of microplankton distribution, and demonstrated the importance of a SCM, co-located with the thermocline, dominated by a single species, the dinoflagellate Ceratium fusus. This species was dominant in the SCM over a wide area of the NW European shelf in the June/July 2015 study period and comprised up to 85% of the SCM biomass. Analysis of similarity and multivariate non-metric multidimensional scaling showed the phytoplankton community of the SCM to be statistically distinct from those of the surface and deep waters. Holography also revealed a fine scale layering of taxa at different levels within the SCM, likely reflecting ecological differences. Some taxa followed the peak abundance of C. fusus, while others reached maximum abundances immediately below or above the C. fusus maximum, suggesting the possible operation of exclusion mechanisms. Additionally, the detection of abundant aggregates located only within and beneath the SCM demonstrates the potential importance of this deep production for the export of carbon to the sea floor. Some predictions of phytoplankton productivity propose a shift to smaller cells in the more stratified oceans of the future resulting in declining production and export. Results presented here, however, contribute to a growing body of evidence that suggests, on the contrary, that key species among the larger celled/colonial, SCM-adapted diatoms and dinoflagellates may instead be selected in stratified conditions, driving increased production and export.

Enhancement of diatom growth and phytoplankton productivity with reduced O2 availability is moderated by rising CO2

Many marine organisms are exposed to decreasing O2 levels due to warming-induced expansion of hypoxic zones and ocean deoxygenation (DeO2). Nevertheless, effects of DeO2 on phytoplankton have been neglected due to technical bottlenecks on examining O2 effects on O2-producing organisms. Here we show that lowered O2 levels increased primary productivity of a coastal phytoplankton assemblage, and enhanced photosynthesis and growth in the coastal diatom Thalassiosira weissflogii. Mechanistically, reduced O2 suppressed mitochondrial respiration and photorespiration of T. weissflogii, but increased the efficiency of their CO2 concentrating mechanisms (CCMs), effective quantum yield and improved light use efficiency, which was apparent under both ambient and elevated CO2 concentrations leading to ocean acidification (OA). While the elevated CO2 treatment partially counteracted the effect of low O2 in terms of CCMs activity, reduced levels of O2 still strongly enhanced phytoplankton primary productivity. This implies that decreased availability of O2 with progressive DeO2 could boost re-oxygenation by diatom-dominated phytoplankton communities, especially in hypoxic areas, with potentially profound consequences for marine ecosystem services in coastal and pelagic oceans.

Aeolian Iron and Its Contribution to Phytoplankton Production in McMurdo Sound, Southwest Ross Sea, Antarctica

<p>Each summer the waters in McMurdo Sound (Lat. 77.5ºS; Long. 165ºE), south-western (SW) Ross Sea encounter vast phytoplankton blooms. This phenomenon is stimulated by the addition of bio-available iron (Fe) to an environment where phytoplankton growth is otherwise Fe-limited. One possible source of such Fe is aeolian sand and dust (ASD) which accumulates on sea ice and is released into the ocean during the summer melt season. The amount of bio-available Fe (i.e. the amount of Fe immedately accessible to phytoplankton) potentially supplied to the ocean by ASD depends on a number of factors including; the ASD flux into the ocean, its particle size distribution and Fe content. However, none of these parameters are well constrained in the SW Ross Sea region and, as a result, the significance of this Fe source in the biogeochemical cycle of phytoplankton growth remains to be quantified. This study focuses on an area (7400 km²) of Southern McMurdo Sound, one of the few areas where direct sampling of ASD that has accumulated on sea ice is possible. To evaluate the flux and solubility of Fe contained in ASD into McMurdo Sound, the mass accumulation rate and particle size of 70 surface snow samples and 3 shallow (3 m) firn cores from the nearby McMurdo Ice Shelf covering the period 2000 - 2008 have been analysed. Selected samples were also measured for total and soluble Fe, Sr and Nd isotopic ratios and mineralogy as a guide to Fe-fertilisation potential and provenance, respectively. Mass and particle size data show an exponential decrease in mass accumulation rate (from 26.00 g m⁻² yr⁻¹ to 0.70 g m⁻² yr⁻¹) and a decrease in modal particle size (from 130 to 69 μm) over a distance of 120 km from Southern McMurdo Sound northwards to Granite Harbour. Both these trends are consistent with ASD being dispersed northwards across the sea ice by southerly storms from an area of the McMurdo Ice Shelf, where submarine freezing and surface ablation have resulted in a surface covered with debris from the sea floor, known as the 'dirty ice' or 'debris bands' (Lat. 77.929ºS; Long. 165.505ºE) in Southern McMurdo Sound. This assertion is further supported by the Sr and Nd isotopic signature of ASD matching local source rocks and the presence of vesicular glass of Southern McMurdo Sound in all samples which also points to the debris bands as the origin of ASD in McMurdo Sound. Bio-available Fe is extremely difficult to quantify hence Fe solubility was used as an approximation in this thesis. Analysis of both total (i.e. particulate and soluble) and the percentage of soluble Fe in the 0.4 - 10 μm dust size fraction (i.e. the fraction most likely to become bio-available) by solution ICP-MS shows a narrow range of values; 3.84 ± 1.99 wt % and 9.42 ± 0.70 % respectively. Combining these values with mass accumulation rate estimates for the particles 0.4 - 10 μm in size, gives an annual soluble Fe flux for the region 500 km² north of the debris bands in McMurdo Sound of 0.55 mg m⁻² yr⁻¹ (9.89 μmol m⁻² yr⁻¹), with spatial variability largely determined by differences in mass accumulation rate. These fluxes are at least an order of magnitude greater than predicted in global dust deposition models for the Southern Ocean and measured in snow samples from East Antarctica. Furthermore, these values exceed the Fe threshold, estimated as 0.2 nM (Boyd and Abraham, 2001), required for phytoplankton growth following the simple dust-biota model of Boyd et al. (2010) and assuming the release of captured ASD in snow is instantaneous. Whilst not constrained in the present study, ASD sourced from the debris bands may be sufficiently widely dispersed, particularly during storm years, to contribute to Fe-fertilisation up to 1200 km from Southern McMurdo Sound. Short, ~10 year long, firn core records of mass accumulation and methylsuphonate concentration, a proxy for phytoplankton productivity, shows a close correspondence between the two during particularly stormy years. Whilst not demonstrating a cause-and-effect relationship, this observation suggests coastal ice cores may contain an important record of the interplay between climate, dust supply, Fe-fertilisation of near shore waters and phytoplankton productivity on decadal and longer timescales.</p>

Aeolian Iron and Its Contribution to Phytoplankton Production in McMurdo Sound, Southwest Ross Sea, Antarctica

<p>Each summer the waters in McMurdo Sound (Lat. 77.5ºS; Long. 165ºE), south-western (SW) Ross Sea encounter vast phytoplankton blooms. This phenomenon is stimulated by the addition of bio-available iron (Fe) to an environment where phytoplankton growth is otherwise Fe-limited. One possible source of such Fe is aeolian sand and dust (ASD) which accumulates on sea ice and is released into the ocean during the summer melt season. The amount of bio-available Fe (i.e. the amount of Fe immedately accessible to phytoplankton) potentially supplied to the ocean by ASD depends on a number of factors including; the ASD flux into the ocean, its particle size distribution and Fe content. However, none of these parameters are well constrained in the SW Ross Sea region and, as a result, the significance of this Fe source in the biogeochemical cycle of phytoplankton growth remains to be quantified. This study focuses on an area (7400 km²) of Southern McMurdo Sound, one of the few areas where direct sampling of ASD that has accumulated on sea ice is possible. To evaluate the flux and solubility of Fe contained in ASD into McMurdo Sound, the mass accumulation rate and particle size of 70 surface snow samples and 3 shallow (3 m) firn cores from the nearby McMurdo Ice Shelf covering the period 2000 - 2008 have been analysed. Selected samples were also measured for total and soluble Fe, Sr and Nd isotopic ratios and mineralogy as a guide to Fe-fertilisation potential and provenance, respectively. Mass and particle size data show an exponential decrease in mass accumulation rate (from 26.00 g m⁻² yr⁻¹ to 0.70 g m⁻² yr⁻¹) and a decrease in modal particle size (from 130 to 69 μm) over a distance of 120 km from Southern McMurdo Sound northwards to Granite Harbour. Both these trends are consistent with ASD being dispersed northwards across the sea ice by southerly storms from an area of the McMurdo Ice Shelf, where submarine freezing and surface ablation have resulted in a surface covered with debris from the sea floor, known as the 'dirty ice' or 'debris bands' (Lat. 77.929ºS; Long. 165.505ºE) in Southern McMurdo Sound. This assertion is further supported by the Sr and Nd isotopic signature of ASD matching local source rocks and the presence of vesicular glass of Southern McMurdo Sound in all samples which also points to the debris bands as the origin of ASD in McMurdo Sound. Bio-available Fe is extremely difficult to quantify hence Fe solubility was used as an approximation in this thesis. Analysis of both total (i.e. particulate and soluble) and the percentage of soluble Fe in the 0.4 - 10 μm dust size fraction (i.e. the fraction most likely to become bio-available) by solution ICP-MS shows a narrow range of values; 3.84 ± 1.99 wt % and 9.42 ± 0.70 % respectively. Combining these values with mass accumulation rate estimates for the particles 0.4 - 10 μm in size, gives an annual soluble Fe flux for the region 500 km² north of the debris bands in McMurdo Sound of 0.55 mg m⁻² yr⁻¹ (9.89 μmol m⁻² yr⁻¹), with spatial variability largely determined by differences in mass accumulation rate. These fluxes are at least an order of magnitude greater than predicted in global dust deposition models for the Southern Ocean and measured in snow samples from East Antarctica. Furthermore, these values exceed the Fe threshold, estimated as 0.2 nM (Boyd and Abraham, 2001), required for phytoplankton growth following the simple dust-biota model of Boyd et al. (2010) and assuming the release of captured ASD in snow is instantaneous. Whilst not constrained in the present study, ASD sourced from the debris bands may be sufficiently widely dispersed, particularly during storm years, to contribute to Fe-fertilisation up to 1200 km from Southern McMurdo Sound. Short, ~10 year long, firn core records of mass accumulation and methylsuphonate concentration, a proxy for phytoplankton productivity, shows a close correspondence between the two during particularly stormy years. Whilst not demonstrating a cause-and-effect relationship, this observation suggests coastal ice cores may contain an important record of the interplay between climate, dust supply, Fe-fertilisation of near shore waters and phytoplankton productivity on decadal and longer timescales.</p>

Exploring the use of compound-specific carbon isotopes as a palaeoproductivity proxy off the coast of Adélie Land, East Antarctica

The Antarctic coastal zone is an area of high primary productivity, particularly within coastal polynyas, where large phytoplankton blooms and drawdown of CO2 occur. Reconstruction of historical primary productivity changes and the associated driving factors could provide baseline insights on the role of these areas as sinks for atmospheric CO2, especially in the context of projected changes in coastal Antarctic sea ice. Here we investigate the potential for using carbon isotopes (δ13C) of fatty acids in marine sediments as a proxy for primary productivity. We use a highly resolved sediment core from off the coast of Adélie Land spanning the last ∼ 400 years and monitor changes in the concentrations and δ13C of fatty acids along with other proxy data from the same core. We discuss the different possible drivers of their variability and argue that C24 fatty acid δ13C predominantly reflects phytoplankton productivity in open-water environments, while C18 fatty acid δ13C reflects productivity in the marginal ice zone. These new proxies have implications for better understanding carbon cycle dynamics in the Antarctica coastal zone in future palaeoclimate studies.

Re-assessing copper and nickel enrichments as paleo-productivity proxies

Copper (Cu) and nickel (Ni) are elements frequently enriched in sedimentary deposits rich in organic matter (OM). In the marine environment, they are mainly supplied to the sediments in association with sedimentary OM. In current environments, a good correlation between the intensity of phytoplankton productivity and the quantities of Cu & Ni transferred to sediments made it possible to establish paleo-productivity calculations based on the contents of ancient sediments in these two metals. The present study is a re-evaluation of the value that can be attributed to these two metals as paleo-productivity proxies. The approach adopted here is based on the examination of a large database already available in the scientific literature. The choice was made to favor the examination of a large amount of data by simple means: comparisons of total organic carbon (TOC) content, enrichment in Cu & Ni (or even other trace metals), and value of the Fe:Al ratio that makes it possible to assess the availability of reactive iron. The basic idea is that the examination of a large number of geological formations thanks to the large database makes it possible to encompass all kinds of paleo-environmental settings, thus comprising an extreme variety of the factors conventionally involved in the mechanisms of accumulation of OM. The aim is to identify strong trends, valid in a large number of paleo-situations, which will have to be carefully taken into account in future detailed paleo-environmental reconstructions. It emerges from this study that, in many cases, Cu and Ni cannot be considered as faithfully reflecting the quantity of OM initially deposited. Several factors acting on the loss of Cu and Ni can be retained, and among them, (1) a rapid loss linked to the decomposition of the OM before the conditions conducive to sulfate-reduction set in; (2) a low abundance of reactive iron which limits the quantity of pyrite liable to form, which significantly hampers Cu & Ni fixation in sediments. If Cu & Ni are not reliably retained in the sediments, that is, proportional to the quantity of OM supplied to the sediment, the paleoenvironmental reconstitutions involving the concentrations of these metals may provide underestimated values of paleoproductivity. An interesting clue is the Fe:Al ratio that makes it possible to quickly know whether the values of the Cu & Ni enrichments are likely to be "abnormally" low.

Relationships Between Northern Hemisphere Teleconnection Patterns and Phytoplankton Productivity in the Neva Estuary (Northeastern Baltic Sea)

Teleconnection patterns can be an important tool for investigating the impact of climate change on biological communities. The aim of the study was, using 2003–2020 data on chlorophyll a concentrations (CHL) and plankton primary production (PP) in midsummer, to determine which of the teleconnection patterns have most pronounced effects on phytoplankton productivity in the estuary located on the border between western and eastern Europe. CHL correlated significantly with the winter values of the North Atlantic Oscillation (NAOw) and Scandinavia (SCANDw) indices, as well as with the values of the annual Polar/Eurasian (POLy) and annual Arctic Oscillation (AOy) indices. PP was significantly correlated with the values of POLy. East Atlantic/Western Russia pattern showed no significant correlation with both phytoplankton indicators. Stepwise multiple linear regressions were performed to determine the most influential indices affecting CHL and PP in the Neva Estuary. POLy, SCANDw, and NAOw appeared to be the main predictors in CHL multiple regression model, while the values of POLy and the July NAO and SCAND values were the main predictors in the PP model. According to our research, the productivity of phytoplankton in the Neva Estuary, located in the most northeastern part of the Baltic Sea, showed a significant relationship with the POL, which determines weather conditions in the northeastern regions of Eurasia. Possible mechanisms of the influence of these teleconnection patterns on phytoplankton productivity are discussed. Using the obtained multi-regression equations and the values of climatic indices, we calculated the values of CHL and PP for 1951–2002 and compared them with the results of field observations. The calculated and measured values of CHL and PP showed a significant increase in phytoplankton productivity in the Neva Estuary in the second half of the 2010s compared to earlier periods. In some years of the 1950s, 1980s, and late 1990s, CHL could also be above average and the low phytoplankton productivity should have been observed in the 1960s–1970s. This indicates a significant contribution of current climate change to fluctuation in phytoplankton productivity observed in recent decades, which should be taken into account when developing measures to protect aquatic ecosystems from eutrophication.

Arctic Ocean stratification set by sea level and freshwater inputs since the last ice age

Salinity-driven density stratification of the upper Arctic Ocean isolates sea-ice cover and cold, nutrient-poor surface waters from underlying warmer, nutrient-rich waters. Recently, stratification has strengthened in the western Arctic but has weakened in the eastern Arctic; it is unknown if these trends will continue. Here we present foraminifera-bound nitrogen isotopes from Arctic Ocean sediments since 35,000 years ago to reconstruct past changes in nutrient sources and the degree of nutrient consumption in surface waters, the latter reflecting stratification. During the last ice age and early deglaciation, the Arctic was dominated by Atlantic-sourced nitrate and incomplete nitrate consumption, indicating weaker stratification. Starting at 11,000 years ago in the western Arctic, there is a clear isotopic signal of Pacific-sourced nitrate and complete nitrate consumption associated with the flooding of the Bering Strait. These changes reveal that the strong stratification of the western Arctic relies on low-salinity inflow through the Bering Strait. In the central Arctic, nitrate consumption was complete during the early Holocene, then declined after 5,000 years ago as summer insolation decreased. This sequence suggests that precipitation and riverine freshwater fluxes control the stratification of the central Arctic Ocean. Based on these findings, ongoing warming will cause strong stratification to expand into the central Arctic, slowing the nutrient supply to surface waters and thus limiting future phytoplankton productivity.

Controlling Factors of Phytoplankton Productivity in Marshes in a Hot Climate with High Seasonal Variation

In this work the Auda marsh, which is part of a system of Iraqi marshes, was sampled to assess the seasonal dynamics and controlling factors of microalgae productivity. The marshes are situated in a hot climate with high seasonal variation near the Arabian Gulf. Physicochemical and biological measurements were taken for water in three areas. Bio-optical models were constructed to describe the primary productivity and chlorophyll-a concentrations in the wet and dry seasons separately and also for the entire area of the Iraqi marshes. The models, as well as almost all measurements, showed high seasonal variation. The mean water temperature was 16 °C in the wet season and 28 °C in the dry season. An almost twofold difference was measured for turbidity and the concentrations of dissolved oxygen and chlorophyll-a for the two seasons. Chlorophyll-a appeared to be a better indicator of ecosystem conditions than primary productivity or biological oxygen demand, according to the results obtained from canonical correlation analysis. Nitrogen or phosphorous did not explain primary productivity or chlorophyll-a to an appreciable extent. Biological variables were related most strongly to water temperature and turbidity, which were the factors most important for controlling phytoplankton productivity in the marshes.