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'.

Reconciling models of primary production and photoacclimation [Invited].

Primary 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 θm, 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 θm 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.

Special Issue on Remote Sensing of Ocean Color: Theory and Applications.

The editorial team are delighted to present this Special Issue of Sensors focused on Remote Sensing of Ocean Color: Theory and Applications. We believe that this is a timely opportunity to showcase current developments across a broad range of topics in ocean color remote sensing (OCRS). Although the field is well-established, in this Special Issue we are able to highlight advances in the applications of the technology, our understanding of the underpinning science, and its relevance in the context of monitoring climate change and engaging public participation.

A Printable Device for Measuring Clarity and Colour in Lake and Nearshore Waters.

Two expanding areas of science and technology are citizen science and three-dimensional (3D) printing. Citizen science has a proven capability to generate reliable data and contribute to unexpected scientific discovery. It can put science into the hands of the citizens, increasing understanding, promoting environmental stewardship, and leading to the production of large databases for use in environmental monitoring. 3D printing has the potential to create cheap, bespoke scientific instruments that have formerly required dedicated facilities to assemble. It can put instrument manufacturing into the hands of any citizen who has access to a 3D printer. In this paper, we present a simple hand-held device designed to measure the Secchi depth and water colour (Forel Ule scale) of lake, estuarine and nearshore regions. The device is manufactured with marine resistant materials (mostly biodegradable) using a 3D printer and basic workshop tools. It is inexpensive to manufacture, lightweight, easy to use, and accessible to a wide range of users. It builds on a long tradition in optical limnology and oceanography, but is modified for ease of operation in smaller water bodies, and from small watercraft and platforms. We provide detailed instructions on how to build the device and highlight examples of its use for scientific education, citizen science, satellite validation of ocean colour data, and low-cost monitoring of water clarity, colour and temperature.

The Influence of Temperature and Community Structure on Light Absorption by Phytoplankton in the North Atlantic.

We present a model that estimates the spectral phytoplankton absorption coefficient ( a p h ( λ ) ) of four phytoplankton groups (picophytoplankton, nanophytoplankton, dinoflagellates, and diatoms) as a function of the total chlorophyll-a concentration (C) and sea surface temperature (SST). Concurrent data on a p h ( λ ) (at 12 visible wavelengths), C and SST, from the surface layer (<20 m depth) of the North Atlantic Ocean, were partitioned into training and independent validation data, the validation data being matched with satellite ocean-colour observations. Model parameters (the chlorophyll-specific phytoplankton absorption coefficients of the four groups) were tuned using the training data and found to compare favourably (in magnitude and shape) with results of earlier studies. Using the independent validation data, the new model was found to retrieve total a p h ( λ ) with a similar performance to two earlier models, using either in situ or satellite data as input. Although more complex, the new model has the advantage of being able to determine a p h ( λ ) for four phytoplankton groups and of incorporating the influence of SST on the composition of the four groups. We integrate the new four-population absorption model into a simple model of ocean colour, to illustrate the influence of changes in SST on phytoplankton community structure, and consequently, the blue-to-green ratio of remote-sensing reflectance. We also present a method of propagating error through the model and illustrate the technique by mapping errors in group-specific a p h ( λ ) using a satellite image. We envisage the model will be useful for ecosystem model validation and assimilation exercises and for investigating the influence of temperature change on ocean colour.

An Ocean-Colour Time Series for Use in Climate Studies: The Experience of the Ocean-Colour Climate Change Initiative (OC-CCI).

Ocean colour is recognised as an Essential Climate Variable (ECV) by the Global Climate Observing System (GCOS); and spectrally-resolved water-leaving radiances (or remote-sensing reflectances) in the visible domain, and chlorophyll-a concentration are identified as required ECV products. Time series of the products at the global scale and at high spatial resolution, derived from ocean-colour data, are key to studying the dynamics of phytoplankton at seasonal and inter-annual scales; their role in marine biogeochemistry; the global carbon cycle; the modulation of how phytoplankton distribute solar-induced heat in the upper layers of the ocean; and the response of the marine ecosystem to climate variability and change. However, generating a long time series of these products from ocean-colour data is not a trivial task: algorithms that are best suited for climate studies have to be selected from a number that are available for atmospheric correction of the satellite signal and for retrieval of chlorophyll-a concentration; since satellites have a finite life span, data from multiple sensors have to be merged to create a single time series, and any uncorrected inter-sensor biases could introduce artefacts in the series, e.g., different sensors monitor radiances at different wavebands such that producing a consistent time series of reflectances is not straightforward. Another requirement is that the products have to be validated against in situ observations. Furthermore, the uncertainties in the products have to be quantified, ideally on a pixel-by-pixel basis, to facilitate applications and interpretations that are consistent with the quality of the data. This paper outlines an approach that was adopted for generating an ocean-colour time series for climate studies, using data from the MERIS (MEdium spectral Resolution Imaging Spectrometer) sensor of the European Space Agency; the SeaWiFS (Sea-viewing Wide-Field-of-view Sensor) and MODIS-Aqua (Moderate-resolution Imaging Spectroradiometer-Aqua) sensors from the National Aeronautics and Space Administration (USA); and VIIRS (Visible and Infrared Imaging Radiometer Suite) from the National Oceanic and Atmospheric Administration (USA). The time series now covers the period from late 1997 to end of 2018. To ensure that the products meet, as well as possible, the requirements of the user community, marine-ecosystem modellers, and remote-sensing scientists were consulted at the outset on their immediate and longer-term requirements as well as on their expectations of ocean-colour data for use in climate research. Taking the user requirements into account, a series of objective criteria were established, against which available algorithms for processing ocean-colour data were evaluated and ranked. The algorithms that performed best with respect to the climate user requirements were selected to process data from the satellite sensors. Remote-sensing reflectance data from MODIS-Aqua, MERIS, and VIIRS were band-shifted to match the wavebands of SeaWiFS. Overlapping data were used to correct for mean biases between sensors at every pixel. The remote-sensing reflectance data derived from the sensors were merged, and the selected in-water algorithm was applied to the merged data to generate maps of chlorophyll concentration, inherent optical properties at SeaWiFS wavelengths, and the diffuse attenuation coefficient at 490 nm. The merged products were validated against in situ observations. The uncertainties established on the basis of comparisons with in situ data were combined with an optical classification of the remote-sensing reflectance data using a fuzzy-logic approach, and were used to generate uncertainties (root mean square difference and bias) for each product at each pixel.

Distribution and antibiotic resistance of vibrio population in an urbanized tropical lake-the Vembanad-in the southwest coast of India.

Among the diverse Vibrio spp. autochthonous to coastal ecosystems, V. cholerae, V. fluvialis, V. vulnificus and V. parahaemolyticus are pathogenic to humans. Increasing sea-surface temperature, sea-level rise and water-related disasters associated with climate change have been shown to influence the proliferation of these bacteria and change their geographic distribution. We investigated the spatio-temporal distribution of Vibrio spp. in a tropical lake for 1 year at a 20-day interval. The abundance of Vibrio spp. was much higher during the south-west monsoon in 2018, when the lake experienced a once-in-a-century flood. The distribution of Vibrio spp. was influenced by salinity (r = 0.3, p < 0.001), phosphate (r = 0.18, p < 0.01) and nitrite (r = 0.16, p < 0.02) in the water. We isolated 470 colonies of Vibrio-like organisms and 341 could be revived further and identified using 16S rRNA gene sequencing. Functional annotations showed that all the 16 Vibrio spp. found in the lake could grow in association with animals. More than 60% of the isolates had multiple antibiotic resistance (MAR) index greater than 0.5. All isolates were resistant to erythromycin and cefepime. The proliferation of multiple antibiotic-resistant Vibrio spp. is a threat to human health. Our observations suggest that the presence of a diverse range of Vibrio spp. is favoured by the low-saline conditions brought about by heavy precipitation. Furthermore, infections caused by contact with Vibrio-contaminated waters may be difficult to cure due to their multiple antibiotic resistances. Therefore, continuous monitoring of bacterial pollution in the lakes is essential, as is the generation of risk maps of vibrio-infested waters to avoid public contact with contaminated waters and associated disease outbreaks.

Analysis of non-pharmaceutical interventions and their impacts on COVID-19 in Kerala.

In the absence of an effective vaccine or drug therapy, non-pharmaceutical interventions are the only option for control of the outbreak of the coronavirus disease 2019, a pandemic with global implications. Each of the over 200 countries affected has followed its own path in dealing with the crisis, making it difficult to evaluate the effectiveness of measures implemented, either individually, or collectively. In this paper we analyse the case of the south Indian state of Kerala, which received much attention in the international media for its actions in containing the spread of the disease in the early months of the pandemic, but later succumbed to a second wave. We use a model to study the trajectory of the disease in the state during the first four months of the outbreak. We then use the model for a retrospective analysis of measures taken to combat the spread of the disease, to evaluate their impact. Because of the differences in the trajectory of the outbreak in Kerala, we argue that it is a model worthy of a place in the discussion on how the world might best handle this and other, future, pandemics.

The Sorcerer II Global Ocean Sampling expedition: northwest Atlantic through eastern tropical Pacific.

The world's oceans contain a complex mixture of micro-organisms that are for the most part, uncharacterized both genetically and biochemically. We report here a metagenomic study of the marine planktonic microbiota in which surface (mostly marine) water samples were analyzed as part of the Sorcerer II Global Ocean Sampling expedition. These samples, collected across a several-thousand km transect from the North Atlantic through the Panama Canal and ending in the South Pacific yielded an extensive dataset consisting of 7.7 million sequencing reads (6.3 billion bp). Though a few major microbial clades dominate the planktonic marine niche, the dataset contains great diversity with 85% of the assembled sequence and 57% of the unassembled data being unique at a 98% sequence identity cutoff. Using the metadata associated with each sample and sequencing library, we developed new comparative genomic and assembly methods. One comparative genomic method, termed "fragment recruitment," addressed questions of genome structure, evolution, and taxonomic or phylogenetic diversity, as well as the biochemical diversity of genes and gene families. A second method, termed "extreme assembly," made possible the assembly and reconstruction of large segments of abundant but clearly nonclonal organisms. Within all abundant populations analyzed, we found extensive intra-ribotype diversity in several forms: (1) extensive sequence variation within orthologous regions throughout a given genome; despite coverage of individual ribotypes approaching 500-fold, most individual sequencing reads are unique; (2) numerous changes in gene content some with direct adaptive implications; and (3) hypervariable genomic islands that are too variable to assemble. The intra-ribotype diversity is organized into genetically isolated populations that have overlapping but independent distributions, implying distinct environmental preference. We present novel methods for measuring the genomic similarity between metagenomic samples and show how they may be grouped into several community types. Specific functional adaptations can be identified both within individual ribotypes and across the entire community, including proteorhodopsin spectral tuning and the presence or absence of the phosphate-binding gene PstS.

Satellite Ocean Colour: Current Status and Future Perspective.

Spectrally resolved water-leaving radiances (ocean colour) and inferred chlorophyll concentration are key to studying phytoplankton dynamics at seasonal and interannual scales, for a better understanding of the role of phytoplankton in marine biogeochemistry; the global carbon cycle; and the response of marine ecosystems to climate variability, change and feedback processes. Ocean colour data also have a critical role in operational observation systems monitoring coastal eutrophication, harmful algal blooms, and sediment plumes. The contiguous ocean-colour record reached 21 years in 2018; however, it is comprised of a number of one-off missions such that creating a consistent time-series of ocean-colour data requires merging of the individual sensors (including MERIS, Aqua-MODIS, SeaWiFS, VIIRS, and OLCI) with differing sensor characteristics, without introducing artefacts. By contrast, the next decade will see consistent observations from operational ocean colour series with sensors of similar design and with a replacement strategy. Also, by 2029 the record will start to be of sufficient duration to discriminate climate change impacts from natural variability, at least in some regions. This paper describes the current status and future prospects in the field of ocean colour focusing on large to medium resolution observations of oceans and coastal seas. It reviews the user requirements in terms of products and uncertainty characteristics and then describes features of current and future satellite ocean-colour sensors, both operational and innovative. The key role of in situ validation and calibration is highlighted as are ground segments that process the data received from the ocean-colour sensors and deliver analysis-ready products to end-users. Example applications of the ocean-colour data are presented, focusing on the climate data record and operational applications including water quality and assimilation into numerical models. Current capacity building and training activities pertinent to ocean colour are described and finally a summary of future perspectives is provided.

Retrieval of phytoplankton size from bio-optical measurements: theory and applications.

The absorption coefficient of a substance distributed as discrete particles in suspension is less than that of the same material dissolved uniformly in a medium-a phenomenon commonly referred to as the flattening effect. The decrease in the absorption coefficient owing to flattening effect depends on the concentration of the absorbing pigment inside the particle, the specific absorption coefficient of the pigment within the particle, and on the diameter of the particle, if the particles are assumed to be spherical. For phytoplankton cells in the ocean, with diameters ranging from less than 1 µm to more than 100 µm, the flattening effect is variable, and sometimes pronounced, as has been well documented in the literature. Here, we demonstrate how the in vivo absorption coefficient of phytoplankton cells per unit concentration of its major pigment, chlorophyll a, can be used to determine the average cell size of the phytoplankton population. Sensitivity analyses are carried out to evaluate the errors in the estimated diameter owing to potential errors in the model assumptions. Cell sizes computed for field samples using the model are compared qualitatively with indirect estimates of size classes derived from high performance liquid chromatography data. Also, the results are compared quantitatively against measurements of cell size in laboratory cultures. The method developed is easy-to-apply as an operational tool for in situ observations, and has the potential for application to remote sensing of ocean colour data.

Water-column stratification governs the community structure of subtropical marine picophytoplankton.

The increase in the areal extent of the subtropical gyres over the past decade has been attributed to a global tendency towards increased water-column stratification. Here, we examine how vertical stratification governs the community structure of the picophytoplankton that dominate these vast marine ecosystems. We analysed phytoplankton community composition in the three Southern Subtropical basins of the Pacific, Indian and Atlantic Oceans using a variety of methods and show that the distributions of picocyanobacteria and photosynthetic picoeukaryotes (PPEs) are strongly correlated with depth and strength of vertical mixing: the changes in community structure occur at various taxonomic levels. In well-mixed waters, PPEs, in particular haptophytes, dominate, whereas in strongly stratified waters, picocyanobacteria of the genus Prochlorococcus are prevalent, regardless of whether the relative contributions to total biomass are assessed in terms of pigment or of carbon. This ecological diochotomy within the picophytoplankton supports the hypothesis that genomic streamlining provides a selective advantage for Prochlorococcus in highly stable, oligotrophic systems, but may restrict their ability to dominate in regions subject to dynamic mixing.

Oceanographic basis of the global surface distribution of Prochlorococcus ecotypes.

By using data collected during a continuous circumnavigation of the Southern Hemisphere, we observed clear patterns in the population-genetic structure of Prochlorococcus, the most abundant photosynthetic organism on Earth, between and within the three Southern Subtropical Gyres. The same mechanisms that were previously invoked to account for the vertical distribution of ecotypes at local scales accounted for the global (horizontal) patterns we observed. Basin-scale and seasonal variations in the structure and strength of vertical stratification provide a basis for understanding large-scale horizontal distribution in genetic and physiological traits of Prochlorococcus, and perhaps of marine microbial communities in general.