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 (ETRRCII, 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 ETRRCII: CO2-assimilation conversion factors and carbon-based primary productivity from FRRF data, without the need for any additional measurements.

We examined the light absorption properties and light-dependent rates of photosynthetic electron transport (ETR RCII ) and 14 C-uptake in phytoplankton of the Canadian Subarctic and Arctic Ocean. Our results reveal high variability in the light-saturated, chlorophyll a-specific rate of 14 C-uptake (P Chla max ; 0.7 to 8.0 mg C mg chl a 21 h 21 ) , and the light-dependant efficiency of 14 C-uptake (a Chla ; 0.01 to 0.09 mg C mg Chl a 21 h 21 [lmol quanta m 22 s 21 ] 21 ). Variability in P Chla max correlated with light availability in the stratified watercolumn, while both P Chla max and a Chla were correlated to the degree of nitrate depletion in the mixed layer. For all samples, we observed that 14 C-uptake rates reached light saturation at lower irradiances than ETR RCII , leading to significant, light-dependent de-coupling of carbon fixation and photosynthetic electron transport. For samples taken from within the mixed layer, we found that the conversion factor from ETR RCII to carbon fixation was strongly correlated to the magnitude of non-photochemical quenching, as derived from fast repetition rate fluorometry (FRRF) measurements. This correlation supports recent findings from the Subarctic Pacific Ocean, and has the potential to improve FRRF-based estimates of phytoplankton carbon fixation. Our observations reveal distinct environmental regulation of phytoplankton light absorption, electron transport and carbon fixation in phytoplankton assemblages within and below the shallow summer mixed layer, and will help to inform more robust predictions of future primary production in a rapidly changing Arctic Ocean.

Abstract. 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 (ETR RCII ), which must be converted into ecologically relevant units of carbon fixation. Understanding sources of variability in the coupling of ETR RCII 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 ETR RCII in the iron-limited NE subarctic Pacific over the course of a diurnal cycle. We show that rates of ETR RCII 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 ETR RCII and carbon fixation (K c /n PSII ). 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 K c /n PSII requires further validation but has the potential to improve estimates of phytoplankton carbon fixation rates from FRRF measurements alone.

Abstract. Active chlorophyll a fluorescence approaches, including fast repetition rate fluorometry (FRRF), have the potential to provide estimates of phytoplankton primary productivity at unprecedented spatial and temporal resolution. FRRF-derived productivity rates are based on estimates of charge separation at PSII (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 coupling ETRRCII and carbon fixation (Φe:C / 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 expression of non-photochemical quenching (NPQ) in the pigment antenna, as derived from FRRF measurements. The observed correlation between NPQ and the conversion factor Φe:C / nPSII has the potential to improve estimates of phytoplankton carbon fixation rates from FRRF measurements alone.

There is an intricate interaction between iron (Fe) and copper (Cu) physiology in diatoms. However, strategies to cope with low Cu are largely unknown. This study unveils the comprehensive restructuring of the photosynthetic apparatus in the diatom Thalassiosira oceanica (CCMP1003) in response to low Cu, at the physiological and proteomic level. The restructuring results in a shift from light harvesting for photochemistry—and ultimately for carbon fixation—to photoprotection, reducing carbon fixation and oxygen evolution. The observed decreases in the physiological parameters Fv/Fm, carbon fixation, and oxygen evolution, concomitant with increases in the antennae absorption cross section (σPSII), non-photochemical quenching (NPQ) and the conversion factor (φe:C/ηPSII) are in agreement with well documented cellular responses to low Fe. However, the underlying proteomic changes due to low Cu are very different from those elicited by low Fe. Low Cu induces a significant four-fold reduction in the Cu-containing photosynthetic electron carrier plastocyanin. The decrease in plastocyanin causes a bottleneck within the photosynthetic electron transport chain (ETC), ultimately leading to substantial stoichiometric changes. Namely, 2-fold reduction in both cytochrome b6f complex (cytb6f) and photosystem II (PSII), no change in the Fe-rich PSI and a 40- and 2-fold increase in proteins potentially involved in detoxification of reactive oxygen species (ferredoxin and ferredoxin:NADP+ reductase, respectively). Furthermore, we identify 48 light harvesting complex (LHC) proteins in the publicly available genome of T. oceanica and provide proteomic evidence for 33 of these. The change in the LHC composition within the antennae in response to low Cu underlines the shift from photochemistry to photoprotection in T. oceanica (CCMP1003). Interestingly, we also reveal very significant intra-specific strain differences. Another strain of T. oceanica (CCMP 1005) requires significantly higher Cu concentrations to sustain both its maximal and minimal growth rate compared to CCMP 1003. Under low Cu, CCMP 1005 decreases its growth rate, cell size, Chla and total protein per cell. We argue that the reduction in protein per cell is the main strategy to decrease its cellular Cu requirement, as none of the other parameters tested are affected. Differences between the two strains, as well as differences between the well documented responses to low Fe and those presented here in response to low Cu are discussed.

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Abstract. Understanding the dynamics of marine phytoplankton productivity requires mechanistic insight into the non-linear coupling of light absorption, photosynthetic electron transport and carbon fixation in response to environmental variability. In the present study, we examined the variability of phytoplankton light absorption characteristics, light-dependent electron transport and 14C-uptake rates over a 48 h period in the coastal subarctic north-east (NE) Pacific. We observed an intricately coordinated response of the different components of the photosynthetic process to diurnal irradiance cycles, which acted to maximize carbon fixation, while simultaneously preventing damage by excess absorbed light energy. In particular, we found diurnal adjustments in pigment ratios, excitation energy transfer to reaction centre II (RCII), the capacity for non-photochemical quenching (NPQ), and the light efficiency (α) and maximum rates (Pmax) of RCII electron transport (ETRRCII) and 14C uptake. Comparison of these results from coastal waters to previous observations in offshore waters of the subarctic NE Pacific provides insight into the effects of iron limitation on the optimization of photosynthesis. Under iron-limited, low-biomass conditions, there was a significant reduction of iron-rich photosynthetic units per chlorophyll a, which was partly offset by higher light absorption and electron transport per photosystem II (PSII). Iron deficiency limited the capacity of phytoplankton to utilize peak midday irradiance for carbon fixation and caused an upregulation of photoprotective mechanisms, including NPQ, and the decoupling of light absorption, electron transport and carbon fixation. Such decoupling resulted in an increased electron requirement (Φe,C) and decreased quantum efficiency (ΦC) of carbon fixation at the iron-limited station. In both coastal and offshore waters, Φe,C and ΦC correlated strongly to NPQ, albeit with a significantly different slope. We discuss the implications of our results for the interpretation of bio-optical data and the parameterization of numerical productivity models, both of which are vital tools in monitoring marine photosynthesis over large temporal and spatial scales.

Abstract. In an experimental assessment of the potential impact of Arctic Ocean acidification on seasonal phytoplankton blooms and associated dimethyl sulfide (DMS) dynamics, we incubated water from Baffin Bay under conditions representing an acidified Arctic Ocean. Using two light regimes simulating under-ice or subsurface chlorophyll maxima (low light; low PAR and no UVB) and ice-free (high light; high PAR + UVA + UVB) conditions, water collected at 38 m was exposed over 9 days to 6 levels of decreasing pH from 8.1 to 7.2. A phytoplankton bloom dominated by the centric diatoms Chaetoceros spp. reaching up to 7.5 µg chlorophyll a L −1 took place in all experimental bags. Total dimethylsulfoniopropionate (DMSP T ) and DMS concentrations reached 155 and 19 nmol L −1 , respectively. The sharp increase in DMSP T and DMS concentrations coincided with the exhaustion of NO − 3 in most microcosms, suggesting that nutrient stress stimulated DMS(P) synthesis by the diatom community. Under both light regimes, chlorophyll a and DMS concentrations decreased linearly with increasing proton concentration at all pH levels tested. Concentrations of DMSP T also decreased but only under high light and over a smaller pH range (from 8.1 to 7.6). In contrast to nanophytoplankton (2-20 µm), pico-phytoplankton (≤ 2 µm) was stimulated by the decreasing pH. We furthermore observed no significant difference between the two light regimes tested in term of chlorophyll a, phytoplankton abundance and taxonomy, and DMSP and DMS net concentrations. These results show that ocean acidification could significantly decrease the algal biomass and inhibit DMS production during the seasonal phytoplankton bloom in the Arctic, with possible consequences for the regional climate.