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Global Ocean Phytoplankton

Marine phytoplankton are responsible for roughly half the net primary production (NPP) on Earth, fixing atmospheric CO2 into food that fuels global ocean ecosystems and drives the ocean's biogeochemical cycles. Phytoplankton growth is highly sensitive to variations in ocean physical properties, such as upper ocean stratification and light availability within this mixed layer. Satellite ocean color sensors, such as the Sea-viewing Wide Field-of-view Sensor (SeaWiFS; McClain 2009) and Moderate Resolution Imaging Spectroradiometer (MODIS; Esaias 1998), provide observations of sufficient frequency and geographic coverage to globally monitor physically-driven changes in phytoplankton distributions. In practice, ocean color sensors retrieve the spectral distribution of visible solar radiation reflected upward from beneath the ocean surface, which can then be related to changes in the photosynthetic phytoplankton pigment, chlorophyll- a (Chla; measured in mg m-3). Here, global Chla data for 2013 are evaluated within the context of the 16-year continuous record provided through the combined observations of SeaWiFS (1997-2010) and MODIS on Aqua (MODISA; 2002-present). Ocean color measurements from the recently launched Visible and Infrared Imaging Radiometer Suite (VIIRS; 2011-present) are also considered, but results suggest that the temporal calibration of the VIIRS sensor is not yet sufficiently stable for quantitative global change studies. All MODISA (version 2013.1), SeaWiFS (version 2010.0), and VIIRS (version 2013.1) data presented here were produced by NASA using consistent Chla algorithms.

phytoplankton↗

Impact of Growth Phase, Pigment Adaptation, and Climate Change Conditions on the Cellular Pigment and Carbon Content of Fifty‐One Phytoplankton Isolates

Owing to their importance in aquatic ecosystems, the demand for models that estimate phytoplankton biomass and community composition in the global ocean has increased over the last decade. Moreover, the impacts of climate change, including elevated carbon dioxide (CO2), increased stratification, and warmer sea surface temperatures, will likely shape phytoplankton community composition in the global ocean. Chemotaxonomic methods are useful for modeling phytoplankton community composition from marker pigments normalized to chlorophyll a (Chl a). However, photosynthetic pigments, particularly Chl a, are sensitive to nutrient and light conditions. Cellular carbon is less sensitive, so using carbon biomass instead may provide an alternative approach. To this end, cellular pigment and carbon concentrations were measured in 51 strains of globally relevant, cultured phytoplankton. Pigment-to-Chl a and pigment-to-carbon ratios were computed for each strain. For 25 strains, measurements were taken during two growth phases. While some differences between growth phases were observed, they did not exceed within class differences. Multiple strains of Amphidinium carterae, Ditylum brightwellii and Heterosigma akashiwo were measured to determine whether time in culture influenced pigment and carbon composition. No appreciable trends in cellular pigment or carbon content were observed. Lastly, the potential impact of climate change conditions on the pigment ratios was assessed using a multistressor experiment that included increased mean light, temperature, and elevated pCO2 on three species: Thalassiosira oceanica, Ostreococcus lucimarinus, and Synechococcus. The largest differences were observed in the pigment-to-carbon ratios, while the marker pigments largely covaried with Chl a. The implications of these observations to chemotaxonomic applications are discussed.

Pigments↗

Acclimation of Phytoplankton Fe:C Ratios Dampens the Biogeochemical Response to Varying Atmospheric Deposition of Soluble Iron

Abstract Dissolved iron (dFe) plays an important role in regulating marine productivity. In high nutrient, low chlorophyll regions (>33% of the global ocean), iron is the primary growth limiting nutrient, and elsewhere iron can regulate nitrogen fixation by diazotrophs. The link between iron availability and carbon export is strongly dependent on the phytoplankton iron quotas or cellular Fe:C ratios. This ratio varies by more than an order of magnitude in the open ocean and is positively correlated with ambient dFe concentrations in field observations. Representing Fe:C ratios within models is necessary to investigate how ocean carbon cycling will interact with perturbations to iron cycling in a changing climate. The Community Earth System Model ocean component was modified to simulate dynamic, group‐specific, phytoplankton Fe:C that varies as a function of ambient iron concentration. The simulated Fe:C ratios improve the representation of the spatial trends in the observed Fe:C ratios. The acclimation of phytoplankton Fe:C ratios dampens the biogeochemical response to varying atmospheric deposition of soluble iron, compared to a fixed Fe:C ratio. However, varying atmospheric soluble iron supply has first order impacts on global carbon and nitrogen fluxes and on nutrient limitation spatial patterns. Our results suggest that pyrogenic Fe is a significant dFe source that rivals mineral dust inputs in some regions. Changes in dust flux and iron combustion sources (anthropogenic and wildfires) will modify atmospheric Fe inputs in the future. Accounting for dynamic phytoplankton iron quotas is critical for understanding ocean biogeochemistry and projecting its response to variations in atmospheric deposition.

Environmental Sciences & Ecology↗

The role of zinc in the adaptive evolution of polar phytoplankton

Zinc is an essential trace metal for oceanic primary producers with the highest concentrations in polar oceans. However, its role in the biological functioning and adaptive evolution of polar phytoplankton remains enigmatic. Here, we have applied a combination of evolutionary genomics, quantitative proteomics, co-expression analyses and cellular physiology to suggest that model polar phytoplankton species have a higher demand for zinc because of elevated cellular levels of zinc-binding proteins. We propose that adaptive expansion of regulatory zinc-finger protein families, co-expanded and co-expressed zinc-binding proteins families involved in photosynthesis and growth in these microalgal species and their natural communities were identified to be responsible for the higher zinc demand. The expression of their encoding genes in eukaryotic phytoplankton metatranscriptomes from pole-to-pole was identified to correlate not only with dissolved zinc concentrations in the upper ocean but also with temperature, suggesting that environmental conditions of polar oceans are responsible for an increased demand of zinc. Furthermore, these results suggest that zinc plays an important role in supporting photosynthetic growth in eukaryotic polar phytoplankton and that this has been critical for algal colonization of low-temperature polar oceans.

59 BASIC BIOLOGICAL SCIENCES↗

Light-independent phytoplankton degradation and detoxification of methylmercury in water

Phytoplankton serves as a key entry point for the trophic transfer and bioaccumulation of the neurotoxin methylmercury (MeHg) in aquatic food webs. However, it is unclear whether and how phytoplankton itself may degrade and metabolize MeHg in the dark. Here, using several strains of the freshwater alga Chlorella vulgaris, the marine diatom Chaetoceros gracilis and two cyanobacteria (or blue-green algae), we report a light-independent pathway of MeHg degradation in water by phytoplankton, rather than its associated bacteria. About 36–85% of MeHg could be degraded intracellularly to inorganic Hg(II) and/or Hg(0) via dark reactions. Furthermore, endogenic reactive oxygen species, particularly singlet oxygen, were identified as the main driver of MeHg demethylation. Given the increasing incidence of algal blooms in lakes and marine systems globally, these findings underscore the potential roles of phytoplankton demethylation and detoxification of MeHg in aquatic ecosystems and call for improved modelling and assessment of MeHg bioaccumulation and environmental risks.

59 BASIC BIOLOGICAL SCIENCES↗

Laboratory tank studies of a single species of phytoplankton using a remote sensing fluorosensor

Phytoplankton were grown in the laboratory for the purpose of testing a remote fluorosensor. The fluorosensor uses a unique four-wavelength dye laser system to excite phytoplankton bearing chlorophyll and to measure the chlorophyll fluorescence generated by this excitation. Six different species were tested, one at a time, and each was grown two to four times. Fluorescence measured by the fluorosensor provides good quantitative measurement of chlorophyll concentrations for all species tested while the cultures were in log phase growth. Fluorescene cross section ratios obtained in the single species tank tests support the hypothesis that the shape of the fluorescence cross section curve remains constant with the species (differences in fluorescence cross section ratios are a basis for determining composition of phytoplankton according to color group when a multiwavelength source of excitation is used. Linear relationships exist between extracted chlorophyll concentration and fluorescence measured by the remote fluorosensor during the log phase growth of phytoplankton cultures tested.

Brown, C. A., Jr.↗

Interpretation of an index of phytoplankton population composition calculated from Remote Airborne Fluorsensor (RAF) data

The calculation of indices of phytoplankton population composition from chlorophyll a fluorescence at 685 nm excited by narrow band light at 454 and 539 nm is discussed. The ratio of the fluorescence excited by light of these two wavelengths is a function of the distribution of the phytoplankton between two color groups, designated the golden-brown and the green. The golden-brown group consists of those species which have the highly photosynthetically active carotenoid-chlorophyll-a-protein complexes, i.e. members of the classes Bacillariophyceae, diatoms Dinophyceae, dinoflagellates, and some members of the class Prymnesiophyceae. The green color group consists those species of phytoplankton which apparently lack those complexes, i.e. members of the classes Chlorophyceae, Euglenophyceae, Prasinophyceae, Eustigmatophyceae, Xanthophyceae, and a few members of the Prymnesiophyceae. A few species of phytoplankton appear to have intermediate characteristics, and would apparently belong to neither group. Most of these species are members of the class Cryptophyceae. The composition index for this class is examined in detail.

Farmer, F. H.↗

Use of ordination and classification procedures to evaluate phytoplankton communities during Superflux II

Cluster analysis and an ordination procedure were performed on two data matrices to investigate real and environmental spatial relationships. Multiple regression analysis was used to relate the measured environmental variables to the phytoplankton community changes. Qualitative type phytoplankton data proved to be less structured in both of these spaces, relative to the biomass data. The salinity gradients of the northern transects covaried significantly with the phytoplankton association changes. In the southern transects the light variable was most important in explaining the variance in the ordination axes. These data suggest the close relationships between phytoplankton community changes and the physical hydrology of the area.

Rutledge, C. K.↗

Light response of phytoplankton in the South Atlantic Ocean: Interpretation of observations and application to remote sensing

A simplified, nonspectral derivation of a classical theory in plant physiology is presented and used to derive an absorption-based primary productivity algorithm. Field observations from a meridional transect (4 deg N to 42 deg S) in the Atlantic Ocean are then described and interpreted in this theoretical context. The observations include photosynthesis-irradiance curve parameters (alpha and P(sub max)), chlorophyll a and phaeopigment concentration, and estimated phytoplankton absorption coefficients at wavelength = 440 nm (a(sub ph)(440)). Observations near the top (50% I(sub 0)) and bottom (6% I(sub 0)) of the euphotic zone are contrasted. At both light levels, alpha, P(sub max), a(sub ph)(440), and pigment concentration varied similarly along the transect: values were highest at the equator and at the southern end of the transect and lowest in the central South Atlantic. It is concluded that this pattern was related to increased nutrient availability due to equatorial upwelling in the north, and increased wind mixing in the south. At the 50% light level, alpha increased relative to a(sub ph) at the southern end of the transect. This result appears to reflect a large-scale meridional (southward) increase in the average quantum efficiency of the photosynthetic units of the phytoplankton. A correlation analysis of the data reveals that at the 50% light level, variations in P(sub max) were more closely related to a(sub ph)(440) than chlorophyll concentration and that phytoplankton absorption explains 90% of the variability in P(sub max). In theory, this shows that the ratio of the average quantum efficiency of the photosynthetic units of the phytoplankton to the product of their average absorption cross section and turnover time is relatively constant. This result is used to simplify the absorption-based primary productivity algorithm derived previously. The feasibility of using this model to estimate production rate from satellite ocean color observations is discussed. It is concluded that an absorption-based algorithm should provide more accurate production rate estimates than one based upon chlorophyll (pigment) concentration.

Hood, Raleigh R.↗

In situ phytoplankton absorption, fluorescence emission, and particulate backscattering spectra determined from reflectance

An inverse model was developed to extract the absortion and scattering (elastic and inelastic) properties of oceanic constituents from surface spectral reflectance measurements. In particular, phytoplankton spectral absorption coefficients, solar-stimulated chlorophyll a fluorescence spectra, and particle backscattering spectra were modeled. The model was tested on 35 reflectance spectra obtained from irradiance measurements in optically diverse ocean waters (0.07 to 25.35 mg/cu m range in surface chlorophyll a concentrations). The universality of the model was demonstrated by the accurate estimation of the spectral phytoplankton absorption coefficents over a range of 3 orders of magnitude (rho = 0.94 at 500 nm). Under most oceanic conditions (chlorophyll a less than 3 mg/cu m) the percent difference between measured and modeled phytoplankton absorption coefficents was less than 35%. Spectral variations in measured phytoplankton absorption spectra were well predicted by the inverse model. Modeled volume fluorescence was weakly correlated with measured chl a; fluorescence quantum yield varied from 0.008 to 0.09 as a function of environment and incident irradiance. Modeled particle backscattering coefficients were linearly related to total particle cross section over a twentyfold range in backscattering coefficents (rho = 0.996, n = 12).

Roesler, Collin S.↗

Large Scale Variability of Phytoplankton Blooms in the Arctic and Peripheral Seas: Relationships with Sea Ice, Temperature, Clouds, and Wind

Spatially detailed satellite data of mean color, sea ice concentration, surface temperature, clouds, and wind have been analyzed to quantify and study the large scale regional and temporal variability of phytoplankton blooms in the Arctic and peripheral seas from 1998 to 2002. In the Arctic basin, phytoplankton chlorophyll displays a large symmetry with the Eastern Arctic having about fivefold higher concentrations than those of the Western Arctic. Large monthly and yearly variability is also observed in the peripheral seas with the largest blooms occurring in the Bering Sea, Sea of Okhotsk, and the Barents Sea during spring. There is large interannual and seasonal variability in biomass with average chlorophyll concentrations in 2002 and 2001 being higher than earlier years in spring and summer. The seasonality in the latitudinal distribution of blooms is also very different such that the North Atlantic is usually most expansive in spring while the North Pacific is more extensive in autumn. Environmental factors that influence phytoplankton growth were examined, and results show relatively high negative correlation with sea ice retreat and strong positive correlation with temperature in early spring. Plankton growth, as indicated by biomass accumulation, in the Arctic and subarctic increases up to a threshold surface temperature of about 276-277 degree K (3-4 degree C) beyond which the concentrations start to decrease suggesting an optimal temperature or nutrient depletion. The correlation with clouds is significant in some areas but negligible in other areas, while the correlations with wind speed and its components are generally weak. The effects of clouds and winds are less predictable with weekly climatologies because of unknown effects of averaging variable and intermittent physical forcing (e.g. over storm event scales with mixing and upwelling of nutrients) and the time scales of acclimation by the phytoplankton.

Comiso, Josefino C.↗

From Surface Chlorophyll a to Phytoplankton Community Composition in Oceanic Waters

The objective of the present study is to examine the potential of using the near-surface total chlorophyll a concentration (C(sub surf)), as it can be derived from ocean color observation, to infer the column-integrated and the vertical distribution of the phytoplanktonic biomass, both in a quantitative way and in a qualitative way (z.e., in terms of community structure). Within this context, a large HPLC (High Performance Liquid Chromatography) pigment database has been analyzed. It includes 2419 vertical pigment profiles, all sampled in Case-1 waters with various trophic states. The relationshps between C(sub surf) and the total chlorophyll alpha vertical distribution, as previously derived by Morel and Berthon, are fully confirmed, as the present results coincide with the previous ones. This agreement allows to go further, namely to examine the possibility of extracting relationships between C(sub surf) and the vertical composition of the algal assemblages. Thanks to the detailed pigment composition available from HPLC measurements, the contribution of three size classes (micro-, nano-, and pico-phytoplankton) to the local total chlorophyll a concentration can be assessed. Corroborating previous findings (e.g., large species dominate in eutrophc environments, whereas tiny phytoplankton prevail in oligotrophic zones), the results lead to a statistically based parameterization. The predictive skill of this parameterization is successfully tested on a separate data set. With such a tool, the vertical total chlorophyll a profiles associated with each size class can be inferred from the sole knowledge of C(sub surf). By combining this tool with satellite ocean color data, it becomes conceivable to quantify on a global scale the phytoplankton biomass associated with each of the three size classes.

Uitz, Julia↗

Phytoplankton-Fluorescence-Lifetime Vertical Profiler

A battery-operated optoelectronic instrument is designed to be lowered into the ocean to measure the intensity and lifetime of fluorescence of chlorophyll A in marine phytoplankton as a function of depth from 0 to 300 m. Fluorescence lifetimes are especially useful as robust measures of photosynthetic productivity of phytoplankton and of physical and chemical mechanisms that affect photosynthesis. The knowledge of photosynthesis in phytoplankton gained by use of this and related instruments is expected to contribute to understanding of global processes that control the time-varying fluxes of carbon and associated biogenic elements in the ocean. The concentration of chlorophyll in the ocean presents a major detection challenge because in order to obtain accurate values of photosynthetic parameters, the intensity of light used to excite fluorescence must be kept very low so as not to disturb the photosynthetic system. Several innovations in fluorometric instrumentation were made in order to make it possible to reach the required low detection limit. These innovations include a highly efficient optical assembly with an integrated flow-through sample interface, and a high-gain, low-noise electronic detection subsystem. The instrument also incorporates means for self-calibration during operation, and electronic hardware and software for control, acquisition and analysis of data, and communications. The electronic circuitry is highly miniaturized and designed to minimize power demand. The instrument is housed in a package that can withstand the water pressure at the maximum depth of 300 m. A light-emitting diode excites fluorescence in the sample flow cell, which is placed at one focal point of an ellipsoidal reflector. A photomultiplier tube is placed at the other focal point. This optical arrangement enables highly efficient collection of fluorescence emitted over all polar directions. Fluorescence lifetime is measured indirectly, by use of a technique based on the same principle as the one described in "Fluorometer for Analysis of Photosynthesis in Phytoplankton" (SSC-00110), NASA Tech Briefs, Vol. 24, No. 1 (November 2000), page 79. The excitation is modulated at a frequency of 70 MHz, and the phase shift between the excitation light and the emitted fluorescence is measured by a detection method in which the 70 MHz signal is down-converted to a 400 Hz signal. The fluorescence lifetime can be computed from the known relationship among the fluorescence lifetime, phase shift, and modulation frequency

Fernandez, Salvador M.↗

State of Climate 2011 - Global Ocean Phytoplankton

Phytoplankton photosynthesis in the sun lit upper layer of the global ocean is the overwhelmingly dominant source of organic matter that fuels marine ecosystems. Phytoplankton contribute roughly half of the global (land and ocean) net primary production (NPP; gross photosynthesis minus plant respiration) and phytoplankton carbon fixation is the primary conduit through which atmospheric CO2 concentrations interact with the ocean s carbon cycle. Phytoplankton productivity depends on the availability of sunlight, macronutrients (e.g., nitrogen, phosphorous), and micronutrients (e.g., iron), and thus is sensitive to climate-driven changes in the delivery of these resources to the euphotic zone

Siegel, D. A.↗

Discriminating Phytoplankton Functional Types (PFTs) in the Coastal Ocean Using the Inversion Algorithm Phydotax and Airborne Imaging Spectrometer Data

There is a need in the Biological Oceanography community to discriminate among phytoplankton groups within the bulk chlorophyll pool to understand energy flow through ecosystems, to track the fate of carbon in the ocean, and to detect and monitor-for harmful algal blooms (HABs). The ocean color community has responded to this demand with the development of phytoplankton functional type (PFT) discrimination algorithms. These PFT algorithms fall into one of three categories depending on the science application: size-based, biogeochemical function, and taxonomy. The new PFT algorithm Phytoplankton Detection with Optics (PHYDOTax) is an inversion algorithm that discriminates taxon-specific biomass to differentiate among six taxa found in the California Current System: diatoms, dinoflagellates, haptophytes, chlorophytes, cryptophytes, and cyanophytes. PHYDOTax was developed and validated in Monterey Bay, CA for the high resolution imaging spectrometer, Spectroscopic Aerial Mapping System with On-board Navigation (SAMSON - 3.5 nm resolution). PHYDOTax exploits the high spectral resolution of an imaging spectrometer and the improved spatial resolution that airborne data provides for coastal areas. The objective of this study was to apply PHYDOTax to a relatively lower resolution imaging spectrometer to test the algorithm's sensitivity to atmospheric correction, to evaluate capability with other sensors, and to determine if down-sampling spectral resolution would degrade its ability to discriminate among phytoplankton taxa. This study is a part of the larger Hyperspectral Infrared Imager (HyspIRI) airborne simulation campaign which is collecting Airborne Visible/Infrared Imaging Spectrometer (AVIRIS) imagery aboard NASA's ER-2 aircraft during three seasons in each of two years over terrestrial and marine targets in California. Our aquatic component seeks to develop and test algorithms to retrieve water quality properties (e.g. HABs and river plumes) in both marine and in-land water bodies. Results presented are from the 10 April 2013 overflight of the Monterey Bay region and focus primarily on the first objective - sensitivity to atmospheric correction. On-going and future work will continue to evaluate if PHYDOTax can be applied to historical (SeaWiFS and MERIS), existing (MODIS, VIIRS, and HICO), and future (PACE, GEO-CAPE, and HyspIRI) satellite sensors. Demonstration of cross-platform continuity may aid in calibration and validation efforts of these sensors.

Biological Oceanography↗

Using Bio-Optics to Reveal Phytoplankton Physiology from a Wirewalker Autonomous Platform

Rapid, wave-powered profiling of bio-optical properties from an autonomous Wirewalker platform provides useful insights into phytoplankton physiology, including the patterns of diel growth, phytoplankton mortality, nonphotochemical quenching of chlorophyll a fluorescence, and natural (sun-induced) fluorescence of mixed communities. Methods are proposed to quantify each of these processes. Such autonomous measurements of phytoplankton physiological rates and responses open up new possibilities for studying phytoplankton in situ, over longer periods, and under a broader range of environmental conditions.

Omand, M. M.↗

Radiative Transfer Modeling of Phytoplankton Fluorescence Quenching Processes

We report the first radiative transfer model that is able to simulate phytoplankton fluorescence with both photo chemical and non-photo chemical quenching included. The fluorescence source term in the inelastic radiative transfer equation is proportional to both the quantum yield and scalar irradiance at excitation wavelengths. The photo chemical and non photo chemical quenching processes change the quantum yield based on the photosynthetic active radiation. A sensitivity study was performed to demonstrate the dependence of the fluorescence signal on chlorophyll a concentration, aerosol optical depths and solar zenith angles. This work enables us to better model the phytoplankton fluorescence, which can be used in the design of new space-based sensors that can provide sufficient sensitivity to detect the phytoplankton fluorescence signal. It could also lead to more accurate remote sensing algorithms for the study of phytoplankton physiology.

Inelastic scattering↗

Remote Sensing of Phytoplankton Pigments

Pigments, as a vital part of phytoplankton, act as the light harvesters and protectors in the process of photosynthesis. Historically, most of the previous studies have been focused on chlorophyll a, the primary light harvesting pigment. With the advances in technologies, especially High-Performance Liquid Chromatography (HPLC)and satellite ocean color remote sensing, recent studies promote the importance of the accessory pigments within the phytoplankton. In this chapter, we will overview the technology advances in phytoplankton pigment identification, the history of ocean color remote sensing and its application in retrieving phytoplankton pigments, and the existing challenges and opportunities for future studies in this field.

Guoqing Wang↗