Remote sensing of phytoplankton community composition in the northern Benguela upwelling system

Marine phytoplankton in the northern Benguela upwelling system (nBUS) serve as a food and energy source fuelling marine food webs at higher trophic levels and thereby support a lucrative fisheries industry that sustain local economies in Namibia. Microscopic and chemotaxonomic analyses are among the most commonly used techniques for routine phytoplankton community analysis and monitoring. However, traditional in situ sampling methods have a limited spatiotemporal coverage. Satellite observations far surpass traditional discrete ocean sampling methods in their ability to provide data at broad spatial scales over a range of temporal resolution over decadal time periods. Recognition of phytoplankton ecological and functional differences has compelled advancements in satellite observations over the past decades to go beyond chlorophyll-a (Chl-a) as a proxy for phytoplankton biomass to distinguish phytoplankton taxa from space. In this study, a multispectral remote sensing approach is presented for detection of dominant phytoplankton groups frequently observed in the nBUS. Here, we use a large microscopic dataset of phytoplankton community structure and the Moderate Resolution Imaging Spectroradiometer of aqua satellite match-ups to relate spectral characteristics of in water constituents to dominance of specific phytoplankton groups. The normalised fluorescence line height, red-near infrared as well as the green/green spectral band-ratios were assigned to the dominant phytoplankton groups using statistical thresholds. The ocean colour remote sensing algorithm presented here is the first to identify phytoplankton functional types in the nBUS with far-reaching potential for mapping the phenology of phytoplankton groups on unprecedented spatial and temporal scales towards advanced ecosystem understanding and environmental monitoring

Projected poleward migration of the Southern Ocean CO2 sink region under high emissions

The Southern Ocean is a major region of ocean carbon uptake, but its future changes remain uncertain under climate change. Here we show the projected shift in the Southern Ocean CO2 sink using a suite of Earth System Models, revealing changes in the mechanism, position and seasonality of the carbon uptake. The region of dominant CO2 uptake shifts from the Subtropical to the Antarctic region under the high-emission scenario. The warming-driven sea-ice melt, increased ocean stratification, mixed layer shoaling, and a weaker vertical carbon gradient is projected to together reduce the winter de-gassing in the future, which will trigger the switch from mixing-driven outgassing to solubility-driven uptake in the Antarctic region during the winter season. The future Southern Ocean carbon sink will be poleward-shifted, operating in a hybrid mode between biologically-driven summertime and solubility-driven wintertime uptake with further amplification of biologically-driven uptake due to the increasing Revelle Factor

Multidecadal trend of increasing iron stress in Southern Ocean phytoplankton.

Southern Ocean primary productivity is principally controlled by adjustments in light and iron limitation, but the spatial and temporal determinants of iron availability, accessibility, and demand are poorly constrained, which hinders accurate long-term projections. We present a multidecadal record of phytoplankton photophysiology between 1996 and 2022 from historical in situ datasets collected by Biogeochemical Argo (BGC-Argo) floats and ship-based platforms. We find a significant multidecadal trend in irradiance-normalized nonphotochemical quenching due to increasing iron stress, with concomitant declines in regional net primary production. The observed trend of increasing iron stress results from changing Southern Ocean mixed-layer physics as well as complex biological and chemical feedback that is indicative of important ongoing changes to the Southern Ocean carbon cycle

A seasonal transition in biological carbon pump efficiency in the northern Scotia Sea, Southern Ocean

The biological carbon pump (BCP) contributes to the oceanic CO2 sink by transferring particulate organic carbon (POC) into the deep ocean. The magnitude and efficiency of the BCP is likely to vary on timescales of days to seasons, however characterising this variability from shipboard observations is challenging. High resolution, sustained observations of primary production and particle fluxes by autonomous vehicles offer the potential to fill this knowledge gap. Here we present a 4 month, daily, 1 m vertical resolution glider dataset, collected in the high productivity bloom, downstream of South Georgia, Southern Ocean. The dataset reveals substantial temporal variability in primary production, POC flux and attenuation. During the pre-bloom peak phase we find high export efficiency, implying minimal heterotrophic POC consumption, i.e. productivity is decoupled from upper ocean remineralisation processes. As the bloom progresses from its peak through its declining phase, export flux decreases, but transfer efficiency within the upper 100 m of the mesopelagic increases. Conversely, transfer efficiency in the lower mesopelagic decreases in the post-bloom phase, implying that the flux attenuation processes operating in the upper and lower mesopelagic are effectively decoupled. This finding underscores an important limitation of using a single parameter, such as Martin's ‘b’, to characterise POC flux attenuation in a given location or season. Frequent pulses of export flux are observed throughout the deployment, indicating decoupling between primary production and the processes driving export of material from the upper ocean. The mechanisms underlying the observed seasonal changes in BCP magnitude and efficiency are unclear, as temperature and oxygen concentration changed minimally, although the nature of the sinking particles changed substantially as the bloom progressed. Our results highlight the difficulty of capturing temporal variability and episodic flux events with traditional shipboard observations, which affects our conceptual understanding of the BCP. The increasing use of autonomous vehicles to observe particle fluxes will be essential to characterising the temporal variability in magnitude and functioning of the BCP

Absence of photophysiological response to iron addition in autumn phytoplankton in the Antarctic sea-ice zone

he high nutrient–low chlorophyll condition of the Southern Ocean is generally thought to be caused by the low bioavailability of micronutrients, particularly iron, which plays an integral role in phytoplankton photosynthesis. Nevertheless, the Southern Ocean experiences seasonal blooms that generally initiate in austral spring, peak in summer, and extend into autumn. This seasonal increase in primary productivity is typically linked to the seasonal characteristics of nutrient and light supply. To better understand the potential limitations on productivity in the Antarctic sea-ice zone (SIZ), the photophysiological response of phytoplankton to iron addition (2.0 nM FeCl3) was investigated during autumn along the Antarctic coast off Dronning Maud Land. Five short-term (24 h) incubation experiments were conducted around Astrid Ridge (68∘ S) and along a 6∘ E transect, where an autumn bloom was identified in the region of the western SIZ. Surface iron concentrations ranged from 0.27 to 1.39 nM around Astrid Ridge, and 0.56 to 0.63 nM along the 6∘ E transect. Contrary to expectation, the photophysiological response of phytoplankton to iron addition, measured through the photosynthetic efficiency and the absorption cross-section for photosystem II, showed no significant responses. It is thus proposed that since the autumn phytoplankton in the SIZ exhibited a lack of an iron limitation at the time of sampling, the ambient iron concentrations may have been sufficient to fulfil the cellular requirements. This provides new insights into extended iron replete post-bloom conditions in the typically assumed iron deficient high nutrient–low chlorophyll Southern Ocea

Sinking Organic Particles in the Ocean—Flux Estimates From in situ Optical Devices

Optical particle measurements are emerging as an important technique for understanding the ocean carbon cycle, including contributions to estimates of their downward flux, which sequesters carbon dioxide (CO2) in the deep sea. Optical instruments can be used from ships or installed on autonomous platforms, delivering much greater spatial and temporal coverage of particles in the mesopelagic zone of the ocean than traditional techniques, such as sediment traps. Technologies to image particles have advanced greatly over the last two decades, but the quantitative translation of these immense datasets into biogeochemical properties remains a challenge. In particular, advances are needed to enable the optimal translation of imaged objects into carbon content and sinking velocities. In addition, different devices often measure different optical properties, leading to difficulties in comparing results. Here we provide a practical overview of the challenges and potential of using these instruments, as a step toward improvement and expansion of their applications

Southern Ocean phytoplankton dynamics and carbon export: insights from a seasonal cycle approach

Quantifying the strength and efficiency of the Southern Ocean biological carbon pump (BCP) and its response to predicted changes in the Earth's climate is fundamental to our ability to predict long-term changes in the global carbon cycle and, by extension, the impact of continued anthropogenic perturbation of atmospheric CO2. There is little agreement, however, in climate model projections of the sensitivity of the Southern Ocean BCP to climate change, with a lack of consensus in even the direction of predicted change, highlighting a gap in our understanding of a major planetary carbon flux. In this review, we summarize relevant research that highlights the important role of fine-scale dynamics (both temporal and spatial) that link physical forcing mechanisms to biogeochemical responses that impact the characteristics of the seasonal cycle of phytoplankton and by extension the BCP. This approach highlights the potential for integrating autonomous and remote sensing observations of fine scale dynamics to derive regionally optimized biogeochemical parameterizations for Southern Ocean models. Ongoing development in both the observational and modelling fields will generate new insights into Southern Ocean ecosystem function for improved predictions of the sensitivity of the Southern Ocean BCP to climate change. This article is part of a discussion meeting issue 'Heat and carbon uptake in the Southern Ocean: the state of the art and future priorities'

Single-Turnover Variable Chlorophyll Fluorescence as a Tool for Assessing Phytoplankton Photosynthesis and Primary Productivity: Opportunities, Caveats and Recommendations

Phytoplankton photosynthetic physiology can be investigated through single-turnover variable chlorophyll fluorescence (ST-ChlF) approaches, which carry unique potential to autonomously collect data at high spatial and temporal resolution. Over the past decades, significant progress has been made in the development and application of ST-ChlF methods in aquatic ecosystems, and in the interpretation of the resulting observations. At the same time, however, an increasing number of sensor types, sampling protocols, and data processing algorithms have created confusion and uncertainty among potential users, with a growing divergence of practice among different research groups. In this review, we assist the existing and upcoming user community by providing an overview of current approaches and consensus recommendations for the use of ST-ChlF measurements to examine in-situ phytoplankton productivity and photo-physiology. We argue that a consistency of practice and adherence to basic operational and quality control standards is critical to ensuring data inter-comparability. Large datasets of inter-comparable and globally coherent ST-ChlF observations hold the potential to reveal large-scale patterns and trends in phytoplankton photo-physiology, photosynthetic rates and bottom-up controls on primary productivity. As such, they hold great potential to provide invaluable physiological observations on the scales relevant for the development and validation of ecosystem models and remote sensing algorithms

The Southern Ocean mixed layer and its boundary fluxes: Fine-scale observational progress and future research priorities

Interactions between the upper ocean and air-ice-ocean fluxes in the Southern Ocean play a critical role in global climate by impacting the overturning circulation and oceanic heat and carbon uptake. Remote and challenging conditions have led to sparse observational coverage, while ongoing field programmes often fail to collect sufficient information in the right place or at the time-space scales required to constrain the variability occurring in the coupled ocean-atmosphere system. Only within the last 10 years have we been able to directly observe and assess the role of the fine-scale ocean and rapidly evolving atmospheric marine boundary layer on the upper limb of the Southern Ocean's overturning circulation. This review summarizes advances in mechanistic understanding, arising in part from observational programmes using autonomous platforms, of the fine-scale processes (1-100 km, hours-seasons) influencing the Southern Ocean mixed layer and its variability. We also review progress in observing the ocean interior connections and the coupled interactions between the ocean, atmosphere and cryosphere that moderate air-sea fluxes of heat and carbon. Most examples provided are for the ice-free Southern Ocean, while major challenges remain for observing the ice-covered ocean. We attempt to elucidate contemporary research gaps and ongoing/future efforts needed to address them. This article is part of a discussion meeting issue 'Heat and carbon uptake in the Southern Ocean: the state of the art and future priorities'

Using Optical Sensors on Gliders to Estimate Phytoplankton Carbon Concentrations and Chlorophyll-to-Carbon Ratios in the Southern Ocean

One approach to deriving phytoplankton carbon biomass estimates (Cphyto) at appropriate scales is through optical products. This study uses a high-resolution glider data set in the Sub-Antarctic Zone (SAZ) of the Southern Ocean to compare four different methods of deriving Cphyto from particulate backscattering and fluorescence-derived chlorophyll (chl-a). A comparison of the methods showed that at low (<0.5 mg m−3) chlorophyll concentrations (e.g., early spring and at depth), all four methods produced similar estimates of Cphyto, whereas when chlorophyll concentrations were elevated one method derived higher concentrations of Cphyto than the others. The use of methods derived from particulate backscattering rather than fluorescence can account for cellular adjustments in chl-a:Cphyto that are not driven by biomass alone. A comparison of the glider chl-a:Cphyto ratios from the different optical methods with ratios from laboratory cultures and cruise data found that some optical methods of deriving Cphyto performed better in the SAZ than others and that regionally derived methods may be unsuitable for application to the Southern Ocean. A comparison of the glider chl-a:Cphyto ratios with output from a complex biogeochemical model shows that although a ratio of 0.02 mg chl-a mg C−1 is an acceptable mean for SAZ phytoplankton (in spring-summer), the model misrepresents the seasonal cycle (with decreasing ratios from spring to summer and low sub-seasonal variability). As such, it is recommended that models expand their allowance for variable chl-a:Cphyto ratios that not only account for phytoplankton acclimation to low light conditions in spring but also to higher optimal chl-a:Cphyto ratios with increasing growth rates in summer