Multi-vehicle adaptive 3D mapping for targeted ocean sampling.

Expanding spatial presentation from two-dimensional profile transects to three-dimensional ocean mapping is key for a better understanding of ocean processes. Phytoplankton distributions can be highly patchy and the accurate identification of these patches with the context, variability, and uncertainty of measurements on relevant scales is difficult to achieve. Traditional sampling methods, such as plankton nets, water samplers and in-situ vertical sensors, provide a snapshot and often miss the fine-scale horizontal and temporal variability. Here, we show how two autonomous underwater vehicles measured, adapted to, and reported real-time chlorophyll a measurements, giving insights into the spatiotemporal distribution of phytoplankton biomass and patchiness. To gain the maximum available information within their sensing scope, the vehicles moved in an adaptive fashion, looking for the regions of the highest predicted chlorophyll a concentration, the greatest uncertainty, and the least possibility of collision with other underwater vehicles and ships. The vehicles collaborated by exchanging data with each other and operators via satellite, using a common segmentation of the area to maximize information exchange over the limited bandwidth of the satellite. Importantly, the use of multiple autonomous underwater vehicles reporting real-time data combined with targeted sampling can provide better match with sampling towards understanding of plankton patchiness and ocean processes.

Use of an uncrewed surface vehicle and near infrared hyperspectral imaging for sampling and analysis of aquatic microplastics.

Data on MP in aquatic environments have low resolution in space and time. Scaling up sampling and increasing analysis throughput are the main bottlenecks. We combined two approaches: an uncrewed surface vehicle (USV) and near infrared hyperspectral imaging (NIR-HSI) for sampling and analysis of MP > 300 µm. We collected 35 water samples over 4 d in a coastal area. Samples were analyzed using NIR-HSI and Fourier transform infrared spectroscopy (FTIR). Spiked samples were used to determine recovery. We conclude that using a USV can mitigate issues of traditional trawls like scalability, repeatability, and contamination. NIR-HSI detects more polyethylene but less polypropylene than FTIR analysis and reduces analysis time significantly. Highly variable concentrations were found at both sampling locations, with mean MP concentration of 0.28 and 0.01 MP m-3 for location A and B respectively. USV sampling in tandem with NIR-HSI is an effective analytical pipeline for MP monitoring.

Development and calibration of a high dynamic range and autonomous ocean-light instrument to measure sub-surface profiles in ice-covered waters.

The optical chain and logger (OptiCAL) is an autonomous ice-tethered observatory equipped with multiple light sensors for mapping the variation of light with depth. We describe the instrument and present an ensemble calibration for downwelling irradiance E P A R in [µm o l m -2 s -1]. Results from a long-term deployment in the Arctic Ocean demonstrate that the OptiCAL can cover the high dynamic range of under-ice light levels from July to November and produce realistic values in terms of magnitude when compared to modeled surface irradiance. Transient features of raised light levels at specific depths associated with nearby leads in the ice underline the importance of depth-resolved light measurements.

Photophysiologically active green, red, and brown macroalgae living in the Arctic Polar Night.

Arctic macroalgae species have developed different growth strategies to survive extreme seasonal changes in irradiance in polar regions. We compared photophysiological parameters such as the light saturation parameter (Ek) and pigment composition of green, red, and brown macroalgae collected in January (Polar Night) and October 2020 (end of the light season). Macroalgae in January appeared healthier (morphologically) and had longer lamina (new growth) than those in October. EK values for red, and brown algae were higher with lower maximum quantum yield of PS II fluorescence (Fv/Fm) in January versus October. Furthermore, in January, new tissues in kelp species had higher EK than the older tissue. Higher EK and lower Fv/Fm during the Polar Night indicates that the photosynthetic apparatus is active but slow. Furthermore, we discuss Chlorophyll (Chl) a emission spectra under blue and green excitation light to determine the ratio of Chl a in photosystem II (PS II) vs photosystem I (PS I). Absorbance spectra of P. palmata was used to interpret the emission spectra. The observed spectral shifts in the absorbance and reflectance spectra of different macroalgae is discussed. Photophysiological methods provide health information complementary to future mapping and monitoring of macroalgae. These results reveal that macroalgae grow new tissue in darkness.

Spectral and RGB analysis of the light climate and its ecological impacts using an all-sky camera system in the Arctic.

The ArcLight observatory provides an hourly continuous time series of all-sky images providing light climate data (intensity, spectral composition, and photoperiod) from the Arctic (Svalbard at 79°N). Until recently, no complete annual time series of light climate relevant for biological processes has been provided from the high Arctic because of insufficient sensitivity of commercial light sensors during the Polar Night. The ArcLight set up is unique, as it provides both all-sky images and the corresponding integrated spectral irradiance in the visible part of the solar electromagnetic spectrum (E P A R ). Here we present a further development providing hourly diel-annual dynamics from 2020 of the irradiance partitioned into the red, green, and blue parts of the solar spectrum and illustrate their relation to weather conditions, and sun and moon trajectories. We show that there is variation between the RGB proportions of irradiance throughout the year, with the blue part of the spectrum showing the greatest variation, which is dependent on weather conditions (i.e., cloud cover). We further provide an example of the biological impact of these spectral variations in the light climate using in vivo Chl a-specific absorption coefficients of diatoms (mean of six low light acclimated northern-Arctic bloom-forming species) to model total algal light absorption (AQ t o t a l ) and the corresponding fraction of quanta used by Photosystem II (AQPSII) (O 2 production) in RGB bands and the potential impacts on the photoreceptor response, suggesting periods where repair and maintenance functions dominate activity in the absence of appreciable levels of red or green light. The method used here can be applied to light climate data and spectral response data worldwide to give localized ecological models of AQ.

Contrasting phytoplankton-zooplankton distributions observed through autonomous platforms, in-situ optical sensors and discrete sampling.

Plankton distributions are remarkably 'patchy' in the ocean. In this study, we investigated the contrasting phytoplankton-zooplankton distributions in relation to wind mixing events in waters around a biodiversity-rich island (Runde) located off the western coast of Norway. We used adaptive sampling from AUV and shipboard profiles of in-situ phytoplankton photo-physiology and particle identification (copepods, fecal pellets and the dinoflagellate Tripos spp.) and quantification using optical and imaging sensors. Additionally, traditional seawater and net sampling were collected for nutrient and in-vitro chlorophyll a concentrations and phytoplankton and meso-zooplankton abundances. Persistent strong wind conditions (~5 days) disrupted the stratification in offshore regions, while stratification and a subsurface chlorophyll maximum (SCM) were observed above the base of the mixed layer depth (MLD ~30 m) in inshore waters. Contrasting phytoplankton and zooplankton abundances were observed between inshore (with the presence of a SCM) and offshore waters (without the presence of a SCM). At the SCM, phytoplankton abundances (Tripos spp., the diatom Proboscia alata and other flagellates) were half (average of 200 cell L-1) of those observed offshore. On the contrary, meso-zooplankton counts were ~6× higher (732 ind m-3 for Calanus spp.) inshore (where a SCM was observed) compared to offshore areas. In parallel, fecal pellets and ammonium concentrations were high (>1000 ind m-3 for the upper 20 m) at the SCM, suggesting that the shallow mixed layer might have increased encounter rates and promoted strong grazing pressure. Low nutrient concentrations (< 1µM for nitrate) were found below the MLD (60 m) in offshore waters, suggesting that mixing and nutrient availability likely boosted phytoplankton abundances. The size of the absorption cross-section (σPII') and yield of photosystem II photochemistry under ambient light (φPII') changed according to depth, while the depth-related electron flow (JPII) was similar between regions, suggesting a high degree of community plasticity to changes in the light regime. Our results emphasize the importance of using multiple instrumentation, in addition to traditional seawater and net sampling for a holistic understanding of plankton distributions.

Photophysiological cycles in Arctic krill are entrained by weak midday twilight during the Polar Night.

Light plays a fundamental role in the ecology of organisms in nearly all habitats on Earth and is central for processes such as vision and the entrainment of the circadian clock. The poles represent extreme light regimes with an annual light cycle including periods of Midnight Sun and Polar Night. The Arctic Ocean extends to the North Pole, and marine light extremes reach their maximum extent in this habitat. During the Polar Night, traditional definitions of day and night and seasonal photoperiod become irrelevant since there are only "twilight" periods defined by the sun's elevation below the horizon at midday; we term this "midday twilight." Here, we characterize light across a latitudinal gradient (76.5° N to 81° N) during Polar Night in January. Our light measurements demonstrate that the classical solar diel light cycle dominant at lower latitudes is modulated during Arctic Polar Night by lunar and auroral components. We therefore question whether this particular ambient light environment is relevant to behavioral and visual processes. We reveal from acoustic field observations that the zooplankton community is undergoing diel vertical migration (DVM) behavior. Furthermore, using electroretinogram (ERG) recording under constant darkness, we show that the main migratory species, Arctic krill (Thysanoessa inermis) show endogenous increases in visual sensitivity during the subjective night. This change in sensitivity is comparable to that under exogenous dim light acclimations, although differences in speed of vision suggest separate mechanisms. We conclude that the extremely weak midday twilight experienced by krill at high latitudes during the darkest parts of the year has physiological and ecological relevance.

All-sky camera system providing high temporal resolution annual time series of irradiance in the Arctic.

The ArcLight observatory provides hourly continuous time series of light regime data (intensity, spectral composition, and photoperiod) from the Arctic, Svalbard at 79° N. Until now, no complete annual time series of biologically relevant light has been provided from the high Arctic due to insufficient sensitivity of commercial light sensors during the Polar Night. We describe a camera system providing all-sky images and the corresponding integrated spectral irradiance (EPAR) in energy or quanta units, throughout a complete annual cycle. We present hourly-diel-annual dynamics from 2017 to 2020 of irradiance and its relation to weather conditions, sun and moon trajectories.

Pelagic organisms avoid white, blue, and red artificial light from scientific instruments.

In situ observations of pelagic fish and zooplankton with optical instruments usually rely on external light sources. However, artificial light may attract or repulse marine organisms, which results in biased measurements. It is often assumed that most pelagic organisms do not perceive the red part of the visible spectrum and that red light can be used for underwater optical measurements of biological processes. Using hull-mounted echosounders above an acoustic probe or a baited video camera, each equipped with light sources of different colours (white, blue and red), we demonstrate that pelagic organisms in Arctic and temperate regions strongly avoid artificial light, including visible red light (575-700 nm), from instruments lowered in the water column. The density of organisms decreased by up to 99% when exposed to artificial light and the distance of avoidance varied from 23 to 94 m from the light source, depending on colours, irradiance levels and, possibly, species communities. We conclude that observations from optical and acoustic instruments, including baited cameras, using light sources with broad spectral composition in the 400-700 nm wavelengths do not capture the real state of the ecosystem and that they cannot be used alone for reliable abundance estimates or behavioural studies.

In situ recordings of large gelatinous spheres from NE Atlantic, and the first genetic confirmation of egg mass of Illex coindetii (Vérany, 1839) (Cephalopoda, Mollusca).

In total, 90 gelatinous spheres, averaging one meter in diameter, have been recorded from ~ 1985 to 2019 from the NE Atlantic Ocean, including the Mediterranean Sea, using citizen science. More than 50% had a dark streak through center. They were recorded from the surface to ~ 60-70 m depth, mainly neutrally buoyant, in temperatures between 8 and 24°C. Lack of tissue samples has until now, prohibited confirmation of species. However, in 2019 scuba divers secured four tissue samples from the Norwegian coast. In the present study, DNA analysis using COI confirms species identity as the ommastrephid broadtail shortfin squid Illex coindetii (Vérany, 1839); these are the first confirmed records from the wild. Squid embryos at different stages were found in different egg masses: (1) recently fertilized eggs (stage ~ 3), (2) organogenesis (stages ~ 17-19 and ~ 23), and (3) developed embryo (stage ~ 30). Without tissue samples from each and every record for DNA corroboration we cannot be certain that all spherical egg masses are conspecific, or that the remaining 86 observed spheres belong to Illex coindetii. However, due to similar morphology and size of these spheres, relative to the four spheres with DNA analysis, we suspect that many of them were made by I. coindetii.

Artificial light during the polar night disrupts Arctic fish and zooplankton behaviour down to 200 m depth.

For organisms that remain active in one of the last undisturbed and pristine dark environments on the planet-the Arctic Polar Night-the moon, stars and aurora borealis may provide important cues to guide distribution and behaviours, including predator-prey interactions. With a changing climate and increased human activities in the Arctic, such natural light sources will in many places be masked by the much stronger illumination from artificial light. Here we show that normal working-light from a ship may disrupt fish and zooplankton behaviour down to at least 200 m depth across an area of >0.125 km2 around the ship. Both the quantitative and qualitative nature of the disturbance differed between the examined regions. We conclude that biological surveys in the dark from illuminated ships may introduce biases on biological sampling, bioacoustic surveys, and possibly stock assessments of commercial and non-commercial species.

Sea surface microlayer and elemental composition in phaeo-, chloro-, and rhodophytes in winter and spring.

Trace metal or rare earth element (REE) content of marine macroalgae are underexamined and there is a great need for further understanding since macroalgae are used for food and may also be bioindicators of environmental changes. This study, by using high resolution inductive coupled plasma spectrometer in a clean laboratory (class 1000), investigates the trace metal and REE concentrations and composition in the youngest tissue of various species within three algal classes in the Trondheimsfjord, Norway, comprising phaeo-, chloro-, and rhodophytes in winter (February) and Spring (May) 2013, with the main focus on phaeophytes. The macroalgae were found in a clear zonation pattern as a function of depth. A significant difference in element concentration and composition was found between the six phaeophyte species along with a significant difference between winter and spring tissue. A zonation depth trend in algal tissue element concentration was also found for the phaeophytes, where the algal species located in both extreme ends (upper vs. lower littoral zone) obtained a lower element concentration than the algae located in the middle part of the littoral zone. This trend seems to result from different algal contact with the metal-rich sea surface microlayer. The chlorophytes had 5-27 times higher concentration of REE and lead (Pb) than the two other algal classes. Results indicate that the rhodo- and chlorophytes are better accumulators than the phaeophytes for several trace metals and REE.

Toward adaptive robotic sampling of phytoplankton in the coastal ocean.

Currents, wind, bathymetry, and freshwater runoff are some of the factors that make coastal waters heterogeneous, patchy, and scientifically interesting-where it is challenging to resolve the spatiotemporal variation within the water column. We present methods and results from field experiments using an autonomous underwater vehicle (AUV) with embedded algorithms that focus sampling on features in three dimensions. This was achieved by combining Gaussian process (GP) modeling with onboard robotic autonomy, allowing volumetric measurements to be made at fine scales. Special focus was given to the patchiness of phytoplankton biomass, measured as chlorophyll a (Chla), an important factor for understanding biogeochemical processes, such as primary productivity, in the coastal ocean. During multiple field tests in Runde, Norway, the method was successfully used to identify, map, and track the subsurface chlorophyll a maxima (SCM). Results show that the algorithm was able to estimate the SCM volumetrically, enabling the AUV to track the maximum concentration depth within the volume. These data were subsequently verified and supplemented with remote sensing, time series from a buoy and ship-based measurements from a fast repetition rate fluorometer (FRRf), particle imaging systems, as well as discrete water samples, covering both the large and small scales of the microbial community shaped by coastal dynamics. By bringing together diverse methods from statistics, autonomous control, imaging, and oceanography, the work offers an interdisciplinary perspective in robotic observation of our changing oceans.

Underwater hyperspectral imaging as an in situ taxonomic tool for deep-sea megafauna.

Identification of benthic megafauna is commonly based on analysis of physical samples or imagery acquired by cameras mounted on underwater platforms. Physical collection of samples is difficult, particularly from the deep sea, and identification of taxonomic morphotypes from imagery depends on resolution and investigator experience. Here, we show how an Underwater Hyperspectral Imager (UHI) can be used as an alternative in situ taxonomic tool for benthic megafauna. A UHI provides a much higher spectral resolution than standard RGB imagery, allowing marine organisms to be identified based on specific optical fingerprints. A set of reference spectra from identified organisms is established and supervised classification performed to identify benthic megafauna semi-autonomously. The UHI data provide an increased detection rate for small megafauna difficult to resolve in standard RGB imagery. In addition, seafloor anomalies with distinct spectral signatures are also detectable. In the region investigated, sediment anomalies (spectral reflectance minimum at ~675 nm) unclear in RGB imagery were indicative of chlorophyll a on the seafloor. Underwater hyperspectral imaging therefore has a great potential in seafloor habitat mapping and monitoring, with areas of application ranging from shallow coastal areas to the deep sea.

Underwater hyperspectral imaging: a new tool for marine archaeology.

By applying an underwater hyperspectral imager (UHI) to a selection of archaeological artifacts, we have found spectral signatures that are representative of materials likely to be present at wreck sites. By successfully using the signatures to classify a subset of said artifacts placed on the seabed at 61 m depth, we demonstrate that it is possible to detect archaeological objects of interest in UHI data acquired by a remotely operated vehicle. Correct UHI classification of rust and glass bottles in situ on a historical wreck site further supports the viability of the method for marine archaeological applications.

Use of an Autonomous Surface Vehicle reveals small-scale diel vertical migrations of zooplankton and susceptibility to light pollution under low solar irradiance.

Light is a major cue for nearly all life on Earth. However, most of our knowledge concerning the importance of light is based on organisms' response to light during daytime, including the dusk and dawn phase. When it is dark, light is most often considered as pollution, with increasing appreciation of its negative ecological effects. Using an Autonomous Surface Vehicle fitted with a hyperspectral irradiance sensor and an acoustic profiler, we detected and quantified the behavior of zooplankton in an unpolluted light environment in the high Arctic polar night and compared the results with that from a light-polluted environment close to our research vessels. First, in environments free of light pollution, the zooplankton community is intimately connected to the ambient light regime and performs synchronized diel vertical migrations in the upper 30 m despite the sun never rising above the horizon. Second, the vast majority of the pelagic community exhibits a strong light-escape response in the presence of artificial light, observed down to 100 m. We conclude that artificial light from traditional sampling platforms affects the zooplankton community to a degree where it is impossible to examine its abundance and natural rhythms within the upper 100 m. This study underscores the need to adjust sampling platforms, particularly in dim-light conditions, to capture relevant physical and biological data for ecological studies. It also highlights a previously unchartered susceptibility to light pollution in a region destined to see significant changes in light climate due to a reduced ice cover and an increased anthropogenic activity.

The advective origin of an under-ice spring bloom in the Arctic Ocean using multiple observational platforms.

Under-ice blooms of phytoplankton in the Chukchi Sea have been observed, with strong implications for our understanding of the production regimes in the Arctic Ocean. Using a combination of satellite remote sensing of phytoplankton biomass, in situ observations under sea ice from an autonomous underwater vehicle (AUV), and in vivo photophysiology, we examined the composition, magnitude and origin of a bloom detected beneath the sea ice Northwest of Svalbard (Southern Yermak Plateau) in May 2010. In situ concentration of up to 20 mg chlorophyll a [Chl a] m-3, were dominated by the northern planktonic spring species of diatoms, Thalassiosira nordenskioeldii, T. antarctica var. borealis, Chaetoceros socialis species complex and Fragilariopsis oceanica. These species were also found south of the marginal ice zone (MIZ). Cells in the water column under the sea ice were typically high-light acclimated, with a mean light saturation index (E k ) of 138 µmol photons m-2 s-1 and a ratio between photoprotective carotenoids (PPC) and Chl a (w:w) of 0.2. Remotely sensed data of [Chl a] showed a 32,000 km2 bloom developing south of the MIZ. In effect, our data suggest that the observed under-ice bloom was in fact a bloom developed in open waters south of the ice edge, and that a combination of northward-flowing water masses and southward drifting sea ice effectively positioned the bloom under the sea ice. This have implications for our general understanding of under-ice blooms, suggesting that their origin and connection with open water may be different in different regions of the Arctic.

Leads in Arctic pack ice enable early phytoplankton blooms below snow-covered sea ice.

The Arctic icescape is rapidly transforming from a thicker multiyear ice cover to a thinner and largely seasonal first-year ice cover with significant consequences for Arctic primary production. One critical challenge is to understand how productivity will change within the next decades. Recent studies have reported extensive phytoplankton blooms beneath ponded sea ice during summer, indicating that satellite-based Arctic annual primary production estimates may be significantly underestimated. Here we present a unique time-series of a phytoplankton spring bloom observed beneath snow-covered Arctic pack ice. The bloom, dominated by the haptophyte algae Phaeocystis pouchetii, caused near depletion of the surface nitrate inventory and a decline in dissolved inorganic carbon by 16 ± 6 g C m-2. Ocean circulation characteristics in the area indicated that the bloom developed in situ despite the snow-covered sea ice. Leads in the dynamic ice cover provided added sunlight necessary to initiate and sustain the bloom. Phytoplankton blooms beneath snow-covered ice might become more common and widespread in the future Arctic Ocean with frequent lead formation due to thinner and more dynamic sea ice despite projected increases in high-Arctic snowfall. This could alter productivity, marine food webs and carbon sequestration in the Arctic Ocean.

Bioluminescence as an ecological factor during high Arctic polar night.

Bioluminescence commonly influences pelagic trophic interactions at mesopelagic depths. Here we characterize a vertical gradient in structure of a generally low species diversity bioluminescent community at shallower epipelagic depths during the polar night period in a high Arctic fjord with in situ bathyphotometric sampling. Bioluminescence potential of the community increased with depth to a peak at 80 m. Community composition changed over this range, with an ecotone at 20-40 m where a dinoflagellate-dominated community transitioned to dominance by the copepod Metridia longa. Coincident at this depth was bioluminescence exceeding atmospheric light in the ambient pelagic photon budget, which we term the bioluminescence compensation depth. Collectively, we show a winter bioluminescent community in the high Arctic with vertical structure linked to attenuation of atmospheric light, which has the potential to influence pelagic ecology during the light-limited polar night.

Divinyl chlorophyll a in the marine eukaryotic protist Alexandrium ostenfeldii (Dinophyceae).

Here it is reported the first detection of DV-chl a together with the usual chl a in the marine dinoflagellate Alexandrium ostenfeldii from the Baltic Sea. Growth response and photosynthetic parameters were examined at two irradiances (80 and 240 µmol photons m(-2) s(-1)) and temperatures (15 °C and 19 °C) in a divinylic strain (AOTV-OS20) versus a monovinylic one (AOTV-OS16), using in vivo chl a fluorescence kinetics of PSII to characterize photosynthetic parameters by pulse amplitude modulated fluorescence, (14)C assimilation rates and toxin analyses. The divinylic isolate exhibited slower growth and stronger sensitivity to high irradiance than normal chl a strain. DV-chl a : chl a ratios decreased along time (from 11.3 to < 0.5 after 10 months) and to restore them sub-cloning and selection of strains with highest DV-chl a content was required. A mutation and/or epigenetic changes in the expression of divinyl reductase gene/s in A. ostenfeldii may explain this altered pigment composition. Despite quite severe limitations (reduced fitness and gradual loss of DV-chl a content), the DV-chl a-containing line in A. ostenfeldii could provide a model organism in photosynthetic studies related with chl biosynthesis and evolution.