Size Class Dependent Relationships between Temperature and Phytoplankton Photosynthesis-Irradiance Parameters in the Atlantic Ocean

Over the past decade, a number of methods have been developed to estimate size-class primary production from either in situ phytoplankton pigment data or remotely-sensed data. In this context, the first objective of this study was to compare two methods of estimating size class specific (micro-, nano-, and pico-phytoplankton) photosynthesis-irradiance (PE) parameters from pigment data. The second objective was to analyse the relationship between environmental variables (temperature, nitrate and PAR) and PE parameters in the different size-classes. A large dataset was used of simultaneous measurements of the PE parameters (n = 1,260) and phytoplankton pigment markers (n = 2,326), from 3 different institutes. There were no significant differences in mean PE parameters of the different size classes between the chemotaxonomic method of Uitz et al. (2008) and the pigment markers and carbon-to-Chl a ratios method of Sathyendranath et al. (2009). For both methods, mean maximum photosynthetic rates (PBm ) for micro-phytoplankton were significantly lower than those for pico-phytoplankton and nano-phytoplankton. The mean light limited slope (�B) for nano-phytoplankton were significantly higher than for the other size taxa. For micro-phytoplankton dominated samples identified using the Sathyendranath et al. (2009) method, both PBm and �B exhibited a significant, positive linear relationship with temperature, whereas for pico-phytoplankton the correlation with temperature was negative. Nano-phytoplankton dominated samples showed a positive correlation between PBm and temperature, whereas for �B and the light saturation parameter (Ek) the correlations were not significant. For the Uitz et al. (2008) method, only micro-phytoplankton PBm , pico-phytoplankton �B, nano- and pico-phytoplankton Ek exhibited significant relationships with temperature. The temperature ranges occupied by the size classes derived using these methods differed. The Uitz et al. (2008) method exhibited a wider temperature range compared to those derived from the Sathyendranath et al. (2009) method. The differences arise from the classification of mixed populations. Based on these patterns, we therefore recommend using the Sathyendranath et al. (2009) method to derive micro-phytoplankton PE parameters at sea water temperatures up to 8◦C during monospecific blooms and the Uitz et al. (2008) method to derive PE parameters of mixed populations over the temperature range from 8 to 18◦C. Both methods exhibited similar relationships between pico-phytoplankton PE parameters and temperatures >18◦C

Scratching beneath the surface: a model to predict the vertical distribution of Prochlorococcus using remote sensing.

The unicellular cyanobacterium Prochlorococcus is the most dominant resident of the subtropical gyres, which are considered to be the largest biomes on Earth. In this study, the spatial and temporal variability in the global distribution of Prochlorococcus was estimated in the Atlantic Ocean using an empirical model based on data from 13 Atlantic Meridional Transect cruises. Our model uses satellite-derived sea surface temperature (SST), remote-sensing reflectance at 443 and 488 nm, and the water temperature at a depth of 200 m from Argo data. The model divides the population of Prochlorococcus into two groups: ProI, which dominates under high-light conditions associated with the surface, and ProII, which favors low-light found near the deep chlorophyll maximum. ProI and ProII are then summed to provide vertical profiles of the concentration of Prochlorococcus cells. This model predicts that Prochlorococcus cells contribute 32 Mt of carbon biomass (7.4×1026 cells) to the Atlantic Ocean, concentrated mainly within the subtropical gyres (35%) and areas near the Equatorial Convergence Zone (30%). When projected globally, 3.4×1027 Prochlorococcus cells represent 171 Mt of carbon biomass, with 43% of this global biomass allocated to the upper ocean (0-45 m depth). Annual cell standing stocks were relatively stable between the years 2003 and 2014, and the contribution of the gyres varies seasonally as gyres expand and contract, tracking changes in light and temperature, with lowest cell abundances during the boreal and austral winter (1.4×1013 cells m-2), when surface cell concentrations were highest (9.8×104 cells ml-1), whereas the opposite scenario was observed in spring-summer (2×1013 cells m-2). This model provides a three-dimensional view of the abundance of Prochlorococcus cells, revealing that Prochlorococcus contributes significantly to total phytoplankton biomass in the Atlantic Ocean, and can be applied using either in-situ measurements at the sea surface (r2=0.83) or remote-sensing observables (r2=0.58)

Vertical structure in chlorophyll profiles: influence on primary production in the Arctic Ocean

Subsurface chlorophyll maximum (SCM) layers are prevalent throughout the Arctic Ocean under stratified conditions and are observed both in the wake of retreating sea ice and in thermally stratified waters. The importance of these layers on the overall productivity of Arctic pelagic ecosystems has been a source of debate. In this study, we consider the three principal factors that govern productivity within SCMs: the shape of the chlorophyll profile, the photophysiological characteristics of phytoplankton and the availability of light in the layer. Using the information on the biological and optical parameters describing the vertical structure of chlorophyll, phytoplankton absorption and photosynthesis–irradiance response curves, a spectrally resolved model of primary production is used to identify the set of conditions under which SCMs are important contributors to water-column productivity. Sensitivity analysis revealed systematic errors in the estimation of primary production when the vertical distribution of chlorophyll was not taken into account, with estimates of water-column production using a non-uniform profile being up to 97% higher than those computed using a uniform one. The relative errors were shown to be functions of the parameters describing the shape of the biomass profile and the light available at the SCM to support photosynthesis. Given that SCM productivity is believed to be largely supported by new nutrients, it is likely that the relative contribution of SCMs to new production would be significantly higher than that to gross primary production. We discuss the biogeochemical and ecological implications of these findings and the potential role of new ocean sensors and autonomous underwater vehicles in furthering the study of SCMs in such highly heterogeneous and remote marine ecosystems. This article is part of the theme issue ‘The changing Arctic Ocean: consequences for biological communities, biogeochemical processes and ecosystem functioning'

High photosynthetic rates associated with pico and nanophytoplankton communities and high stratification index in the North West Atlantic

The biological dynamics of pelagic marine ecosystems are strongly influenced by the size structure and ecological succession of phytoplankton, which in turn modifies photosynthetic efficiency. Variability in photosynthetic rates is closely coupled with changes in community structure, but it is difficult to obtain coincident data at high enough resolution to characterise these changes. In this study, we employ hierarchical cluster analysis on chlorophyll-normalised high performance liquid chromatography (HPLC) pigment concentrations from the North West Atlantic, to identify seasonal successional trends amongst phytoplankton populations. Changes in phytoplankton community were also analysed as a function of mean equivalent spherical diameter (MESD) derived from absorption measurements, photosynthetic rates, water-column stratification and temperature. Well-mixed conditions in spring to early summer were associated with populations of large cells containing high concentrations of fucoxanthin, chlorophyll-c1 and chlorophyll-c2 relative to chlorophyll-a (Chl a). As stratification increased over the course of the summer, these cells were replaced by populations dominated by chlorophyll-b, 19'-hexanoyloxyfucoxanthin, 19'-butanoyloxyfucoxanthin and divinyl chlorophyll-a, indicative of small picophytoplankton. As stratification decreased in autumn, MESD and alloxanthin increased, suggesting the presence of cryptophytes. Positive relationships were found between MESD and the quantum yield of photosynthesis (φm) for 7 out of the 8 phytoplankton clusters identified, while negative relationships between mean mixed layer photosynthetically active radiation and φm and the light limited slope of photosynthesis (αB) were observed for 4 clusters, as a result of nutrient limitation and photo-protection. The highest photosynthetic rates were associated with a pico & nanophytoplankton communities, which increased from spring to late summer as stratification intensified. By contrast, diatom communities had the lowest photosynthetic rates throughout the year. These successional patterns in the dominant phytoplankton size-class and phenology support Margalef's mandala in terms of the relationship between turbulence and community structure. The study sheds new light on assemblages dominated by smaller cells, under warm, stratified conditions, having higher photosynthetic efficiencies, which has implications for the carbon flux in the NW Atlantic

Ecophysiological basis of spatiotemporal patterns in picophytoplankton pigments in the global ocean

Information on the intracellular content and functional diversity of phytoplankton pigments can provide valuable insight on the ecophysiological state of primary producers and the flow of energy within aquatic ecosystems. Combined global datasets of analytical flow cytometry (AFC) cell counts and High-Performance Liquid Chromatography (HPLC) pigment concentrations were used to examine vertical and seasonal variability in the ratios of phytoplankton pigments in relation to indices of cellular photoacclimation. Across all open ocean datasets, the weight-to-weight ratio of photoprotective to photosynthetic pigments showed a strong depth dependence that tracked the vertical decline in the relative availability of light. The Bermuda Atlantic Time-series Study (BATS) dataset revealed a general increase in surface values of the relative concentrations of photoprotective carotenoids from the winter-spring phytoplankton communities dominated by low-light acclimated eukaryotic microalgae to the summer and early autumn communities dominated by high-light acclimated picocyanobacteria. In Prochlorococcus-dominated waters, the vertical decline in the relative contribution of photoprotective pigments to total pigment concentration could be attributed in large part to changes in the cellular content of photosynthetic pigments (PSP) rather than photoprotective pigments (PPP), as evidenced by a depth-dependent increase of the intracellular concentration of the divinyl chlorophyll-a (DVChl-a) whilst the intracellular concentration of the PPP zeaxanthin remained relatively uniform with depth. The ability of Prochlorococcus cells to adjust their DVChl-a cell-1 over a large gradient in light intensity was reflected in more highly variable estimates of carbon-to-Chl-a ratio compared to those reported for other phytoplankton groups. This cellular property is likely the combined result of photoacclimatory changes at the cellular level and a shift in dominant ecotypes. Developing a mechanistic understanding of sources of variability in pigmentation of picocyanobacteria is critical if the pigment markers and bio-optical properties of these cells are to be used to map their biogeography and serve as indicators of photoacclimatory state of subtropical phytoplankton communities more broadly. It would also allow better assessment of effects on, and adaptability of phytoplankton communities in the tropical/subtropical ocean due to climate chang

Seasonal cycling of zinc and cobalt in the south-eastern Atlantic along the GEOTRACES GA10 section

Abstract. We report the distributions and stoichiometry of dissolved zinc (dZn) and cobalt (dCo) in sub-tropical and sub-Antarctic waters of the south-eastern Atlantic Ocean during austral spring 2010 and summer 2011/2012. In sub-tropical surface waters, mixed-layer dZn and dCo concentrations during early spring were 1.60 ± 2.58 nM and 30 ± 11 pM, respectively, compared with summer values of 0.14 ± 0.08 nM and 24 ± 6 pM. The elevated spring dZn concentrations resulted from an apparent offshore transport of elevated dZn at depths between 20–55 m, derived from the Agulhas Bank. In contrast, open-ocean sub-Antarctic surface waters displayed largely consistent inter-seasonal mixed-layer dZn and dCo concentrations of 0.10 ± 0.07 nM and 11 ± 5 pM, respectively. Trace metal stoichiometry, calculated from concentration inventories, suggests a greater overall removal for dZn relative to dCo in the upper water column of the south-eastern Atlantic, with inter-seasonally decreasing dZn / dCo inventory ratios of 19–5 and 13–7 mol mol−1 for sub-tropical surface water and sub-Antarctic surface water, respectively. In this paper, we investigate how the seasonal influences of external input and phytoplankton succession may relate to the distribution of dZn and dCo and variation in dZn / dCo stoichiometry across these two distinct ecological regimes in the south-eastern Atlantic. </jats:p

A global perspective on marine photosynthetic picoeukaryote community structure

A central goal in ecology is to understand the factors affecting the temporal dynamics and spatial distribution of microorganisms and the underlying processes causing differences in community structure and composition. However, little is known in this respect for photosynthetic picoeukaryotes (PPEs), algae that are now recognised as major players in marine CO2 fixation. Here, we analysed dot blot hybridisation and cloning–sequencing data, using the plastid-encoded 16S rRNA gene, from seven research cruises that encompassed all four ocean biomes. We provide insights into global abundance, α- and β-diversity distribution and the environmental factors shaping PPE community structure and composition. At the class level, the most commonly encountered PPEs were Prymnesiophyceae and Chrysophyceae. These taxa displayed complementary distribution patterns, with peak abundances of Prymnesiophyceae and Chrysophyceae in waters of high (25:1) or low (12:1) nitrogen:phosphorus (N:P) ratio, respectively. Significant differences in phylogenetic composition of PPEs were demonstrated for higher taxonomic levels between ocean basins, using Unifrac analyses of clone library sequence data. Differences in composition were generally greater between basins (interbasins) than within a basin (intrabasin). These differences were primarily linked to taxonomic variation in the composition of Prymnesiophyceae and Prasinophyceae whereas Chrysophyceae were phylogenetically similar in all libraries. These data provide better knowledge of PPE community structure across the world ocean and are crucial in assessing their evolution and contribution to CO2 fixation, especially in the context of global climate change

A conceptual approach to partitioning a vertical profile of phytoplankton biomass into contributions from two communities

This is the final version. Available from American Geophysical Union / Wiley via the DOI in this record. These data were collected and made freely available by the International Argo Program and the national programs that contribute to it (https://argo.ucsd.edu, https://www.ocean-ops.org). The Argo Program is part of the Global Ocean Observing System. All data and code used in the paper are provided openly on a GitHub page (https://github.com/rjbrewin/Two-community-phyto-model). This includes an example Jupyter Notebook Python Script, processing this BGC-Argo float and tuning the models. Details of how to run it without having to install software are provided as Supplementary Material to this manuscript.We describe an approach to partition a vertical profile of chlorophyll-a concentration into contributions from two communities of phytoplankton: one (community 1) that resides principally in the turbulent mixed-layer of the upper ocean and is observable through satellite visible radiometry; the other (community 2) residing below the mixed-layer, in a stably stratified environment, hidden from the eyes of the satellite. The approach is tuned to a time-series of profiles from a Biogeochemical-Argo float in the northern Red Sea, selected as its location transitions from a deep mixed layer in winter (characteristic of vertically well-mixed systems) to a shallow mixed layer in the summer with a deep chlorophyll-a maximum (characteristic of vertically stratified systems). The approach is extended to reproduce profiles of particle backscattering, by deriving the chlorophyll-specific backscattering coefficients of the two communities and a background coefficient assumed to be dominated by non-algal particles in the region. Analysis of the float data reveals contrasting phenology of the two communities, with community 1 blooming in winter and 2 in summer, community 1 negatively correlated with epipelagic stratification, and 2 positively correlated. We observe a dynamic chlorophyll-specific backscattering coefficient for community 1 (stable for community 2), positively correlated with light in the mixed-layer, suggesting seasonal changes in photoacclimation and/or taxonomic composition within community 1. The approach has the potential for monitoring vertical changes in epipelagic biogeography and for combining satellite and ocean robotic data to yield a three-dimensional view of phytoplankton distribution.Medical Research Council (MRC)Simons Foundation (SF)King Abdullah University of Science and Technology (KAUST)European Space Agency (ESA)European Space Agency (ESA

Reconciling models of primary production and photoacclimation

This is the final version. Available on open access from the Optical Society of America via the DOI in this recordPrimary production and photoacclimation models are two important classes of physiological models that find applications in remote sensing of pools and fluxes of carbon associated with phytoplankton in the ocean. They are also key components of ecosystem models designed to study biogeochemical cycles in the ocean. So far, these two classes of models have evolved in parallel, somewhat independently of each other. Here we examine how they are coupled to each other through the intermediary of the photosynthesis–irradiance parameters. We extend the photoacclimation model to accommodate the spectral effects of light penetration in the ocean and the spectral sensitivity of the initial slope of the photosynthesis–irradiance curve, making the photoacclimation model fully compatible with spectrally resolved models of photosynthesis in the ocean. The photoacclimation model contains a parameter , which is the maximum chlorophyll-to-carbon ratio that phytoplankton can attain when available light tends to zero. We explore how size-class-dependent values of could be inferred from field data on chlorophyll and carbon content in phytoplankton, and show that the results are generally consistent with lower bounds estimated from satellite-based primary production calculations. This was accomplished using empirical models linking phytoplankton carbon and chlorophyll concentration, and the range of values obtained in culture measurements. We study the equivalence between different classes of primary production models at the functional level, and show that the availability of a chlorophyll-to-carbon ratio facilitates the translation between these classes. We discuss the importance of the better assignment of parameters in primary production models as an important avenue to reduce model uncertainties and to improve the usefulness of satellite-based primary production calculations in climate research.Simons FoundationEuropean Space AgencyNational Centre for Earth ObservationNational Science Foundatio

Obtaining phytoplankton diversity from ocean color: A scientific roadmap for future development

This is the final version. Available from Frontiers Media via the DOI in this record.To improve our understanding of the role of phytoplankton for marine ecosystems and global biogeochemical cycles, information on the global distribution of major phytoplankton groups is essential. Although algorithms have been developed to assess phytoplankton diversity from space for over two decades, so far the application of these data sets has been limited. This scientific roadmap identifies user needs, summarizes the current state of the art, and pinpoints major gaps in long-term objectives to deliver space-derived phytoplankton diversity data that meets the user requirements. These major gaps in using ocean color to estimate phytoplankton community structure were identified as: (a) the mismatch between satellite, in situ and model data on phytoplankton composition, (b) the lack of quantitative uncertainty estimates provided with satellite data, (c) the spectral limitation of current sensors to enable the full exploitation of backscattered sunlight, and (d) the very limited applicability of satellite algorithms determining phytoplankton composition for regional, especially coastal or inland, waters. Recommendation for actions include but are not limited to: (i) an increased communication and round-robin exercises among and within the related expert groups, (ii) the launching of higher spectrally and spatially resolved sensors, (iii) the development of algorithms that exploit hyperspectral information, and of (iv) techniques to merge and synergistically use the various streams of continuous information on phytoplankton diversity from various satellite sensors' and in situ data to ensure long-term monitoring of phytoplankton composition.ESA SEOM SY-4Sci Synergy projectNAS