The Growth Response of Two Diatom Species to Atmospheric Dust from the Last Glacial Maximum

Relief of iron (Fe) limitation in the surface Southern Ocean has been suggested as one driver of the regular glacial-interglacial cycles in atmospheric carbon dioxide (CO$_2$). The proposed cause is enhanced deposition of Fe-bearing atmospheric dust to the oceans during glacial intervals, with consequent effects on export production and the carbon cycle. However, understanding the role of enhanced atmospheric Fe supply in biogeochemical cycles is limited by knowledge of the fluxes and 'bioavailability' of atmospheric Fe during glacial intervals. Here, we assess the effect of Fe fertilization by dust, dry-extracted from the Last Glacial Maximum portion of the EPICA Dome C Antarctic ice core, on the Antarctic diatom species $\textit{Eucampia antarctica}$ and $\textit{Proboscia inermis}$. Both species showed strong but differing reactions to dust addition. $\textit{E. antarctica}$ increased cell number (3880 vs.786 cells mL$^{-1}$), chlorophyll a (51 vs. 3.9 μg mL$^{-1}$) and particulate organic carbon (POC; 1.68 vs. 0.28 μg mL$^{-1}$) production in response to dust compared to controls. $\textit{P. inermis}$ did not increase cell number in response to dust, but chlorophyll $a$ and POC per cell both strongly increased compared to controls (39 vs. 15 and 2.13 vs. 0.95 ng cell$^{-1}$ respectively). The net result of both responses was a greater production of POC and chlorophyll $a$, as well as decreased Si:C and Si:N incorporation ratios within cells. However, $\textit{E, antarctica}$ decreased silicate uptake for the same nitrate and carbon uptake, while $\textit{P. inermis}$ increased carbon and nitrate uptake for the same silicate uptake. This suggests that nutrient utilization changes in response to Fe addition could be driven by different underlying mechanisms between different diatom species. Enhanced supply of atmospheric dust to the surface ocean during glacial intervals could therefore have driven nutrient-utilization changes which could permit greater carbon fixation for lower silica utilization. Additionally, both species responded more strongly to lower amounts of direct Fe chloride addition than they did to dust, suggesting that not all the Fe released from dust was in a bioavailable form available for uptake by diatoms.Natural Environment Research Council (Studentship), Royal Society (Grant ID: RP120096), German Research Foundation (Grant IDs: HO-4217, BR-3794

Distinct iron cycling in a Southern Ocean eddy

Mesoscale eddies are ubiquitous in the iron-limited Southern Ocean, controlling ocean-atmosphere exchange processes, however their influence on phytoplankton productivity remains unknown. Here we probed the biogeochemical cycling of iron (Fe) in a cold-core eddy. In-eddy surface dissolved Fe (dFe) concentrations and phytoplankton productivity were exceedingly low relative to external waters. In-eddy phytoplankton Fe-to-carbon uptake ratios were elevated 2–6 fold, indicating upregulated intracellular Fe acquisition resulting in a dFe residence time of ~1 day. Heavy dFe isotope values were measured for in-eddy surface waters highlighting extensive trafficking of dFe by cells. Below the euphotic zone, dFe isotope values were lighter and coincident with peaks in recycled nutrients and cell abundance, indicating enhanced microbially-mediated Fe recycling. Our measurements show that the isolated nature of Southern Ocean eddies can produce distinctly different Fe biogeochemistry compared to surrounding waters with cells upregulating iron uptake and using recycling processes to sustain themselves

Seasonal Cycles of Phytoplankton and Net Primary Production from Biogeochemical Argo Float Data in the South-West Pacific Ocean

We present annual cycles of chlorophyll a, phytoplankton carbon, nitrate and oxygen for Subtropical (STW), Subantarctic (SAW), and Subantarctic Mode (SAMW) waters near Aotearoa New Zealand from data collected by two Biogeochemical (BGC) Argo floats. We develop two simple models of depth-integrated net primary production (NPP), tuned against 14C-uptake measurements, to compare with Vertically-Generalised Production Model (VGPM) satellite-based estimates of NPP. One model is the simplest possible, and assumes production is proportional to light multiplied by chlorophyll a concentration. The second model modifies the light response profile to account for photoacclimation. In STW at 30–35°S, enhanced production is initiated in austral autumn when the mixed layer deepens to entrain nutrients into the photic zone. For about half the year, there is substantial production within a deep chlorophyll maximum that sits below the mixed layer. Consequently, depth-integrated NPP is only loosely related to surface biomass as imaged from satellite remote-sensing, and BGC Argo-based model estimates of depth-integrated NPP are about double VGPM estimates. In SAW at 45–55°S, production is initiated when vertical mixing decreases in austral spring. Production is largely within the mixed layer, and depth-integrated phytoplankton biomass and depth-integrated NPP follow surface phytoplankton biomass. Model estimates of depth-integrated NPP based on BGC Argo float profiles are comparable with VGPM estimates for the southern water masses

The ongoing need for rates: can physiology and omics come together to co-design the measurements needed to understand complex ocean biogeochemistry?

The necessity to understand the influence of global ocean change on biota has exposed wide-ranging gaps in our knowledge of the fundamental principles that underpin marine life. Concurrently, physiological research has stagnated, in part driven by the advent and rapid evolution of molecular biological techniques, such that they now influence all lines of enquiry in biological oceanography. This dominance has led to an implicit assumption that physiology is outmoded, and advocacy that ecological and biogeochemical models can be directly informed by omics. However, the main modeling currencies are biological rates and biogeochemical fluxes. Here, we ask: how do we translate the wealth of information on physiological potential from omics-based studies to quantifiable physiological rates and, ultimately, to biogeochemical fluxes? Based on the trajectory of the state-of-the-art in biomedical sciences, along with case-studies from ocean sciences, we conclude that it is unlikely that omics can provide such rates in the coming decade. Thus, while physiological rates will continue to be central to providing projections of global change biology, we must revisit the metrics we rely upon. We advocate for the co-design of a new generation of rate measurements that better link the benefits of omics and physiology

Iron, silicate, and light co-limitation of three Southern Ocean diatom species

The effect of combined iron, silicate, and light co-limitation was investigated in the three diatom species Actinocyclus sp. Ehrenberg, Chaetoceros dichaeta Ehrenberg, and Chaetoceros debilis Cleve, isolated from the Southern Ocean (SO). Growth of all species was co-limited by iron and silicate, reflected in a significant increase in the number of cell divisions compared to the control. Lowest relative Si uptake and drastic frustule malformation was found under iron and silicate co-limitation in C. dichaeta, while Si limitation in general caused cell elongation in both Chaetoceros species. Higher light intensities similar to SO surface conditions showed a negative impact on growth of C. dichaeta and Actinocyclus sp. and no effect on C. debilis. This is in contrast to the assumed light limitation of SO diatoms due to deep wind driven mixing. Our results suggest that growth and species composition of Southern Ocean diatoms is influenced by a sensitive interaction of the abiotic factors, iron, silicate, and light

Alteration of Proteins and Pigments Influence the Function of Photosystem I under Iron Deficiency from Chlamydomonas reinhardtii

BACKGROUND: Iron is an essential micronutrient for all organisms because it is a component of enzyme cofactors that catalyze redox reactions in fundamental metabolic processes. Even though iron is abundant on earth, it is often present in the insoluble ferric [Fe (III)] state, leaving many surface environments Fe-limited. The haploid green alga Chlamydomonas reinhardtii is used as a model organism for studying eukaryotic photosynthesis. This study explores structural and functional changes in PSI-LHCI supercomplexes under Fe deficiency as the eukaryotic photosynthetic apparatus adapts to Fe deficiency. RESULTS: 77K emission spectra and sucrose density gradient data show that PSI and LHCI subunits are affected under iron deficiency conditions. The visible circular dichroism (CD) spectra associated with strongly-coupled chlorophyll dimers increases in intensity. The change in CD signals of pigments originates from the modification of interactions between pigment molecules. Evidence from sucrose gradients and non-denaturing (green) gels indicates that PSI-LHCI levels were reduced after cells were grown for 72 h in Fe-deficient medium. Ultrafast fluorescence spectroscopy suggests that red-shifted pigments in the PSI-LHCI antenna were lost during Fe stress. Further, denaturing gel electrophoresis and immunoblot analysis reveals that levels of the PSI subunits PsaC and PsaD decreased, while PsaE was completely absent after Fe stress. The light harvesting complexes were also susceptible to iron deficiency, with Lhca1 and Lhca9 showing the most dramatic decreases. These changes in the number and composition of PSI-LHCI supercomplexes may be caused by reactive oxygen species, which increase under Fe deficiency conditions. CONCLUSIONS: Fe deficiency induces rapid reduction of the levels of photosynthetic pigments due to a decrease in chlorophyll synthesis. Chlorophyll is important not only as a light-harvesting pigment, but also has a structural role, particularly in the pigment-rich LHCI subunits. The reduced level of chlorophyll molecules inhibits the formation of large PSI-LHCI supercomplexes, further decreasing the photosynthetic efficiency

Trophic status of Chlamydomonas reinhardtii influences the impact of iron deficiency on photosynthesis

To investigate the impact of iron deficiency on bioenergetic pathways in Chlamydomonas, we compared growth rates, iron content, and photosynthetic parameters systematically in acetate versus CO2-grown cells. Acetate-grown cells have, predictably (2-fold) greater abundance of respiration components but also, counter-intuitively, more chlorophyll on a per cell basis. We found that phototrophic cells are less impacted by iron deficiency and this correlates with their higher iron content on a per cell basis, suggesting a greater capacity/ability for iron assimilation in this metabolic state. Phototrophic cells maintain both photosynthetic and respiratory function and their associated Fe-containing proteins in conditions where heterotrophic cells lose photosynthetic capacity and have reduced oxygen evolution activity. Maintenance of NPQ capacity might contribute to protection of the photosynthetic apparatus in iron-limited phototrophic cells. Acetate-grown iron-limited cells maintain high growth rates by suppressing photosynthesis but increasing instead respiration. These cells are also able to maintain a reduced plastoquinone pool