In Situ Estimates of Net Primary Production in the Western North Atlantic With Argo Profiling Floats.

The 14C incubation method for net primary production (NPP) has limited spatial/temporal resolution, while satellite approaches cannot provide direct information at depth. With chlorophyll-a and backscatter measurements from BGC-Argo floats, we quantified year-round NPP in the western North Atlantic Ocean using both the Carbon-based Productivity Model (CbPM) and Photoacclimation Productivity Model (PPM). Comparison with NPP profiles from 14C incubation measurements showed advantages and limitations of both models. CbPM reproduced the magnitude of NPP in most cases. However, in the summer the CbPM-based NPP had a large peak in the subsurface, which was an artifact from the subsurface chlorophyll maximum caused by photoacclimation. PPM avoided the artifacts from photoacclimation, but the magnitude of PPM-derived NPP was smaller than the 14C result. Different NPP distribution patterns along a North-South transect in the Western North Atlantic Ocean were observed, including higher winter NPP/lower summer NPP in the south, timing differences in NPP seasonal phenology, and different NPP depth distribution patterns in the summer months. Using a 6-months record of concurrent oxygen and bio-optical measurements from two Argo floats, we also demonstrated the ability of Argo floats to obtain estimates of the net community production to NPP ratio, ranging from 0.3 in July to -1.0 in December 2016. Our results highlight the utility of float bio-optical profiles and indicate that environmental conditions (e.g., light availability, nutrient supply) are major factors controlling the seasonality and spatial (horizontal and vertical) distributions of NPP in the western North Atlantic Ocean.

Phytoplankton community structuring and succession in a competition-neutral resource landscape.

Phytoplankton community composition and succession affect aquatic food webs and biogeochemistry. Resource competition is commonly viewed as an important governing factor for community structuring and this perception is imbedded in modern ecosystem models. Quantitative consideration of the physical spacing between phytoplankton cells, however, suggests that direct competition for growth-limiting resources is uncommon. Here we describe how phytoplankton size distributions and temporal successions are compatible with a competition-neutral resource landscape. Consideration of phytoplankton-herbivore interactions with proportional feeding size ranges yields small-cell dominated size distributions consistent with observations for stable aquatic environments, whereas predator-prey temporal lags and blooming physiologies shift this distribution to larger mean cell sizes in temporally dynamic environments. We propose a conceptual mandala for understanding phytoplankton community composition where species successional series are initiated by environmental disturbance, guided by the magnitude of these disturbances and nutrient stoichiometry, and terminated with the return toward a 'stable solution'. Our conceptual mandala provides a framework for interpreting and modeling the environmental structuring of natural phytoplankton populations.

Evaluation of diagnostic pigments to estimate phytoplankton size classes.

Phytoplankton accessory pigments are commonly used to estimate phytoplankton size classes, particularly during development and validation of biogeochemical models and satellite ocean color-based algorithms. The diagnostic pigment analysis (DPA) is based on bulk measurements of pigment concentrations and relies on assumptions regarding the presence of specific pigments in different phytoplankton taxonomic groups. Three size classes are defined by the DPA: picoplankton, nanoplankton, and microplankton. Until now, the DPA has not been evaluated against an independent approach that provides phytoplankton size calculated on a per-cell basis. Automated quantitative cell imagery of microplankton and some nanoplankton, used in combination with conventional flow cytometry for enumeration of picoplankton and nanoplankton, provide a novel opportunity to perform an independent evaluation of the DPA. Here, we use a data set from the North Atlantic Ocean that encompasses all seasons and a wide range of chlorophyll concentrations (0.18-5.14 mg m-3). Results show that the DPA overestimates microplankton and picoplankton when compared to cytometry data, and subsequently underestimates the contribution of nanoplankton to total biomass. In contrast to the assumption made by the DPA that the microplankton size class is largely made up of diatoms and dinoflagellates, imaging-in-flow cytometry shows significant presence of diatoms and dinoflagellates in the nanoplankton size class. Additionally, chlorophyll b is commonly attributed solely to picoplankton by the DPA, but Chl b-containing phytoplankton are observed with imaging in both nanoplankton and microplankton size classes. We suggest revisions to the DPA equations and application of uncertainties when calculating size classes from diagnostic pigments.

Global satellite-observed daily vertical migrations of ocean animals.

Every night across the world's oceans, numerous marine animals arrive at the surface of the ocean to feed on plankton after an upward migration of hundreds of metres. Just before sunrise, this migration is reversed and the animals return to their daytime residence in the dark mesopelagic zone (at a depth of 200-1,000 m). This daily excursion, referred to as diel vertical migration (DVM), is thought of primarily as an adaptation to avoid visual predators in the sunlit surface layer1,2 and was first recorded using ship-net hauls nearly 200 years ago3. Nowadays, DVMs are routinely recorded by ship-mounted acoustic systems (for example, acoustic Doppler current profilers). These data show that night-time arrival and departure times are highly conserved across ocean regions4 and that daytime descent depths increase with water clarity4,5, indicating that animals have faster swimming speeds in clearer waters4. However, after decades of acoustic measurements, vast ocean areas remain unsampled and places for which data are available typically provide information for only a few months, resulting in an incomplete understanding of DVMs. Addressing this issue is important, because DVMs have a crucial role in global ocean biogeochemistry. Night-time feeding at the surface and daytime metabolism of this food at depth provide an efficient pathway for carbon and nutrient export6-8. Here we use observations from a satellite-mounted light-detection-and-ranging (lidar) instrument to describe global distributions of an optical signal from DVM animals that arrive in the surface ocean at night. Our findings reveal that these animals generally constitute a greater fraction of total plankton abundance in the clear subtropical gyres, consistent with the idea that the avoidance of visual predators is an important life strategy in these regions. Total DVM biomass, on the other hand, is higher in more productive regions in which the availability of food is increased. Furthermore, the 10-year satellite record reveals significant temporal trends in DVM biomass and correlated variations in DVM biomass and surface productivity. These results provide a detailed view of DVM activities globally and a path for refining the quantification of their biogeochemical importance.

Factors driving the seasonal and hourly variability of sea-spray aerosol number in the North Atlantic.

Four North Atlantic Aerosol and Marine Ecosystems Study (NAAMES) field campaigns from winter 2015 through spring 2018 sampled an extensive set of oceanographic and atmospheric parameters during the annual phytoplankton bloom cycle. This unique dataset provides four seasons of open-ocean observations of wind speed, sea surface temperature (SST), seawater particle attenuation at 660 nm (cp,660, a measure of ocean particulate organic carbon), bacterial production rates, and sea-spray aerosol size distributions and number concentrations (NSSA). The NAAMES measurements show moderate to strong correlations (0.56 < R < 0.70) between NSSA and local wind speeds in the marine boundary layer on hourly timescales, but this relationship weakens in the campaign averages that represent each season, in part because of the reduction in range of wind speed by multiday averaging. NSSA correlates weakly with seawater cp,660 (R = 0.36, P << 0.01), but the correlation with cp,660, is improved (R = 0.51, P < 0.05) for periods of low wind speeds. In addition, NAAMES measurements provide observational dependence of SSA mode diameter (dm) on SST, with dm increasing to larger sizes at higher SST (R = 0.60, P << 0.01) on hourly timescales. These results imply that climate models using bimodal SSA parameterizations to wind speed rather than a single SSA mode that varies with SST may overestimate SSA number concentrations (hence cloud condensation nuclei) by a factor of 4 to 7 and may underestimate SSA scattering (hence direct radiative effects) by a factor of 2 to 5, in addition to overpredicting variability in SSA scattering from wind speed by a factor of 5.

Satellite sensor requirements for monitoring essential biodiversity variables of coastal ecosystems.

The biodiversity and high productivity of coastal terrestrial and aquatic habitats are the foundation for important benefits to human societies around the world. These globally distributed habitats need frequent and broad systematic assessments, but field surveys only cover a small fraction of these areas. Satellite-based sensors can repeatedly record the visible and near-infrared reflectance spectra that contain the absorption, scattering, and fluorescence signatures of functional phytoplankton groups, colored dissolved matter, and particulate matter near the surface ocean, and of biologically structured habitats (floating and emergent vegetation, benthic habitats like coral, seagrass, and algae). These measures can be incorporated into Essential Biodiversity Variables (EBVs), including the distribution, abundance, and traits of groups of species populations, and used to evaluate habitat fragmentation. However, current and planned satellites are not designed to observe the EBVs that change rapidly with extreme tides, salinity, temperatures, storms, pollution, or physical habitat destruction over scales relevant to human activity. Making these observations requires a new generation of satellite sensors able to sample with these combined characteristics: (1) spatial resolution on the order of 30 to 100-m pixels or smaller; (2) spectral resolution on the order of 5 nm in the visible and 10 nm in the short-wave infrared spectrum (or at least two or more bands at 1,030, 1,240, 1,630, 2,125, and/or 2,260 nm) for atmospheric correction and aquatic and vegetation assessments; (3) radiometric quality with signal to noise ratios (SNR) above 800 (relative to signal levels typical of the open ocean), 14-bit digitization, absolute radiometric calibration <2%, relative calibration of 0.2%, polarization sensitivity <1%, high radiometric stability and linearity, and operations designed to minimize sunglint; and (4) temporal resolution of hours to days. We refer to these combined specifications as H4 imaging. Enabling H4 imaging is vital for the conservation and management of global biodiversity and ecosystem services, including food provisioning and water security. An agile satellite in a 3-d repeat low-Earth orbit could sample 30-km swath images of several hundred coastal habitats daily. Nine H4 satellites would provide weekly coverage of global coastal zones. Such satellite constellations are now feasible and are used in various applications.

Light color acclimation is a key process in the global ocean distribution of Synechococcus cyanobacteria.

Marine Synechococcus cyanobacteria are major contributors to global oceanic primary production and exhibit a unique diversity of photosynthetic pigments, allowing them to exploit a wide range of light niches. However, the relationship between pigment content and niche partitioning has remained largely undetermined due to the lack of a single-genetic marker resolving all pigment types (PTs). Here, we developed and employed a robust method based on three distinct marker genes (cpcBA, mpeBA, and mpeW) to estimate the relative abundance of all known Synechococcus PTs from metagenomes. Analysis of the Tara Oceans dataset allowed us to reveal the global distribution of Synechococcus PTs and to define their environmental niches. Green-light specialists (PT 3a) dominated in warm, green equatorial waters, whereas blue-light specialists (PT 3c) were particularly abundant in oligotrophic areas. Type IV chromatic acclimaters (CA4-A/B), which are able to dynamically modify their light absorption properties to maximally absorb green or blue light, were unexpectedly the most abundant PT in our dataset and predominated at depth and high latitudes. We also identified populations in which CA4 might be nonfunctional due to the lack of specific CA4 genes, notably in warm high-nutrient low-chlorophyll areas. Major ecotypes within clades I-IV and CRD1 were preferentially associated with a particular PT, while others exhibited a wide range of PTs. Altogether, this study provides important insights into the ecology of Synechococcus and highlights the complex interactions between vertical phylogeny, pigmentation, and environmental parameters that shape Synechococcus community structure and evolution.

Student's tutorial on bloom hypotheses in the context of phytoplankton annual cycles.

Phytoplankton blooms are elements in repeating annual cycles of phytoplankton biomass and they have significant ecological and biogeochemical consequences. Temporal changes in phytoplankton biomass are governed by complex predator-prey interactions and physically driven variations in upper water column growth conditions (light, nutrient, and temperature). Understanding these dependencies is fundamental to assess future change in bloom frequency, duration, and magnitude and thus represents a quintessential challenge in global change biology. A variety of contrasting hypotheses have emerged in the literature to explain phytoplankton blooms, but over time the basic tenets of these hypotheses have become unclear. Here, we provide a "tutorial" on the development of these concepts and the fundamental elements distinguishing each hypothesis. The intent of this tutorial is to provide a useful background and set of tools for reading the bloom literature and to give some suggestions for future studies. Our tutorial is written for "students" at all stages of their career. We hope it is equally useful and interesting to those with only a cursory interest in blooms as those deeply immersed in the challenge of understanding the temporal dynamics of phytoplankton biomass and predicting its future change.

Resurrecting the ecological underpinnings of ocean plankton blooms.

Nutrient and light conditions control phytoplankton division rates in the surface ocean and, it is commonly believed, dictate when and where high concentrations, or blooms, of plankton occur. Yet after a century of investigation, rates of phytoplankton biomass accumulation show no correlation with cell division rates. Consequently, factors controlling plankton blooms remain highly controversial. In this review, we endorse the view that blooms are not governed by abiotic factors controlling cell division, but rather reflect subtle ecosystem imbalances instigated by climate forcings or food-web shifts. The annual global procession of ocean plankton blooms thus represents a report on the recent history of predator-prey interactions modulated by physical processes that, almost coincidentally, also control surface nutrient inputs.

In situ measurements of phytoplankton fluorescence using low cost electronics.

Chlorophyll a fluorometry has long been used as a method to study phytoplankton in the ocean. In situ fluorometry is used frequently in oceanography to provide depth-resolved estimates of phytoplankton biomass. However, the high price of commercially manufactured in situ fluorometers has made them unavailable to some individuals and institutions. Presented here is an investigation into building an in situ fluorometer using low cost electronics. The goal was to construct an easily reproducible in situ fluorometer from simple and widely available electronic components. The simplicity and modest cost of the sensor makes it valuable to students and professionals alike. Open source sharing of architecture and software will allow students to reconstruct and customize the sensor on a small budget. Research applications that require numerous in situ fluorometers or expendable fluorometers can also benefit from this study. The sensor costs US$150.00 and can be constructed with little to no previous experience. The sensor uses a blue LED to excite chlorophyll a and measures fluorescence using a silicon photodiode. The sensor is controlled by an Arduino microcontroller that also serves as a data logger.

Remote identification of the invasive tunicate Didemnum vexillum using reflectance spectroscopy.

Benthic coverage of the invasive tunicate Didemnum vexillum on Georges Bank is largely unknown. Monitoring of D. vexillum coverage is vital to understanding the impact this invasive species will have on the productive fishing grounds of Georges Bank. Here we investigate using reflectance spectroscopy as a method for remote identification of D. vexillum. Using two different systems, a NightSea Dive-Spec and a combination of LED light sources with a hyperspectral radiometer, we collected in-situ measurements of reflectance from D. vexillum colonies. In comparison to reflectance spectra of other common benthic substrates, D. vexillum appears to have a unique spectral signature between 500 and 600 nm. Measuring the slope of the spectrum between these wavelengths appears to be the most robust method for spectral identification. Using derivative analysis or principal component analysis, the reflectance spectra of D. vexillum can be identified among numerous other spectra of common benthic substrates. An optical system consisting of a radiometer, light source, and camera was deployed on a remotely operated vehicle to test the feasibility of using reflectance to assess D. vexillum coverage. Preliminary results, analyzed here, prove the method to be successful for the areas we surveyed and open the way for its use on large-scale surveys.

Improved irradiances for use in ocean heating, primary production, and photo-oxidation calculations.

Accurate calculation of underwater light is fundamental to predictions of upper-ocean heating, primary production, and photo-oxidation. However, most ocean models simulating these processes do not yet incorporate radiative transfer modules for their light calculations. Such models are often driven by above-surface, broadband, daily averaged irradiance or photosynthetically available radiation (PAR) values obtained from climatology or satellite observations, sometimes without correction for sea-surface reflectance, even though surface reflectance can reduce in-water values by more than 20%. We present factors computed by a radiative transfer code that can be used to convert above-surface values in either energy or quantum units to in-water net irradiance, as needed for calculations of water heating, and to in-water PAR, as needed for calculations of photosynthesis and photo-oxidation.

Evaluation of a compact sensor for backscattering and absorption.

Seawater inherent optical properties (IOPs) are key parameters in a wide range of applications in environmental studies and oceanographic research. In particular, the absorption coefficient (a) is the typical IOP used to obtain the concentration of chlorophyll-a in the water-a critical parameter in biological oceanography studies and the backscattering coefficient (b(b)) is used as a measure of turbidity. In this study, we test a novel instrument concept designed to obtain both the absorption and backscattering coefficients. The instrument would emit a collimated monochromatic light beam into the water retrieving the backscattered light intensity as a function of distance from the center of illumination. We use Monte Carlo modeling of light propagation to create an inversion algorithm that translates the signal from such an instrument into values of a and b(b). Our results, based on simulations spanning the bulk of natural values of seawater IOP combinations, indicate that a 6.2 cm diameter instrument with a radial resolution of 1 cm would be capable of predicting b(b) within less than 13.4% relative difference and a within less than 57% relative difference (for 90% of the inverted a values, the relative errors fall below 29.7%). Additionally, these errors could be further reduced by constraining the inversion algorithm with information from concurrent measurements of other IOPs. Such a compact and relatively simple device could have multiple applications for in situ optical measurements, including a and b(b) retrievals from instrumentation mounted on autonomous underwater vehicles. Furthermore, the same methodology could possibly be used for an out-of-water sensor.

The underwater photic environment of Cape Maclear, Lake Malawi: comparison between rock- and sand-bottom habitats and implications for cichlid fish vision.

Lake Malawi boasts the highest diversity of freshwater fishes in the world. Nearshore sites are categorized according to their bottom substrate, rock or sand, and these habitats host divergent assemblages of cichlid fishes. Sexual selection driven by mate choice in cichlids led to spectacular diversification in male nuptial coloration. This suggests that the spectral radiance contrast of fish, the main determinant of visibility under water, plays a crucial role in cichlid visual communication. This study provides the first detailed description of underwater irradiance, radiance and beam attenuation at selected sites representing two major habitats in Lake Malawi. These quantities are essential for estimating radiance contrast and, thus, the constraints imposed on fish body coloration. Irradiance spectra in the sand habitat were shifted to longer wavelengths compared with those in the rock habitat. Beam attenuation in the sand habitat was higher than in the rock habitat. The effects of water depth, bottom depth and proximity to the lake bottom on radiometric quantities are discussed. The radiance contrast of targets exhibiting diffused and spectrally uniform reflectance depended on habitat type in deep water but not in shallow water. In deep water, radiance contrast of such targets was maximal at long wavelengths in the sand habitat and at short wavelengths in the rock habitat. Thus, to achieve conspicuousness, color patterns of rock- and sand-dwelling cichlids would be restricted to short and long wavelengths, respectively. This study provides a useful platform for the examination of cichlid visual communication.

Climate-driven trends in contemporary ocean productivity.

Contributing roughly half of the biosphere's net primary production (NPP), photosynthesis by oceanic phytoplankton is a vital link in the cycling of carbon between living and inorganic stocks. Each day, more than a hundred million tons of carbon in the form of CO2 are fixed into organic material by these ubiquitous, microscopic plants of the upper ocean, and each day a similar amount of organic carbon is transferred into marine ecosystems by sinking and grazing. The distribution of phytoplankton biomass and NPP is defined by the availability of light and nutrients (nitrogen, phosphate, iron). These growth-limiting factors are in turn regulated by physical processes of ocean circulation, mixed-layer dynamics, upwelling, atmospheric dust deposition, and the solar cycle. Satellite measurements of ocean colour provide a means of quantifying ocean productivity on a global scale and linking its variability to environmental factors. Here we describe global ocean NPP changes detected from space over the past decade. The period is dominated by an initial increase in NPP of 1,930 teragrams of carbon a year (Tg C yr(-1)), followed by a prolonged decrease averaging 190 Tg C yr(-1). These trends are driven by changes occurring in the expansive stratified low-latitude oceans and are tightly coupled to coincident climate variability. This link between the physical environment and ocean biology functions through changes in upper-ocean temperature and stratification, which influence the availability of nutrients for phytoplankton growth. The observed reductions in ocean productivity during the recent post-1999 warming period provide insight on how future climate change can alter marine food webs.

Calibrated near-forward volume scattering function obtained from the LISST particle sizer.

The physical nature of particles, such as size, shape, and composition govern their angular light scattering, which is described by the volume scattering function (VSF). Despite the fact that the VSF is one of the most important inherent optical properties, it has rarely been measured in aquatic environments since no commercial instrument exists to measure the full VSF in the field. The commonly used LISST (Laser In Situ Scattering and Transmissometry) particle sizer (Sequoia Scientific, http://www.sequoiasci.com) measures near-forward angular scattering of a laser source (lambda= 670 nm) at 32 logarithmically-spaced photodetectors arranged between 0.08 and 15 degrees and inverts the data to obtain particle size distribution (PSD). In order to calibrate the LISST to provide the near-forward VSF of unknown particle suspensions, we analyzed the scattering of light by polystyrene bead suspensions of known size distributions and composition, and empirically compared it with the results of Mie theory. This (1) allowed us to obtain a set of instrument specific scaling factors needed to retrieve the magnitude of the VSF and (2) provided validation that the shape of the VSF was appropriately obtained.

Uncertainties of inherent optical properties obtained from semianalytical inversions of ocean color.

We present a method to quantify the uncertainties in the in-water constituent absorption and backscattering coefficients obtained from an inversion of remotely sensed reflectance (rrs). We first find a set of positive inversion solutions within a given uncertainty range around the values of the inverted rrs. The uncertainties of the solutions are then computed based on the statistics of these solutions. We demonstrate the uncertainty calculation algorithm using a specific semianalytic inversion model applied to both a field and a simulated data set. When the associated uncertainties are taken into account, the inverted parameters are generally within the uncertainties of the measured (or simulated) parameters, highlighting the success of the inversion and the method to obtain uncertainties. The specific inversion we use, however, fails to retrieve two spectral parameters within a usable range. The method presented is general and can be applied to all existing semianalytical inversion algorithms.