Seafloor primary production in a changing Arctic Ocean.

Phytoplankton and sea ice algae are traditionally considered to be the main primary producers in the Arctic Ocean. In this Perspective, we explore the importance of benthic primary producers (BPPs) encompassing microalgae, macroalgae, and seagrasses, which represent a poorly quantified source of Arctic marine primary production. Despite scarce observations, models predict that BPPs are widespread, colonizing ~3 million km2 of the extensive Arctic coastal and shelf seas. Using a synthesis of published data and a novel model, we estimate that BPPs currently contribute ~77 Tg C y-1 of primary production to the Arctic, equivalent to ~20 to 35% of annual phytoplankton production. Macroalgae contribute ~43 Tg C y-1, seagrasses contribute ~23 Tg C y-1, and microalgae-dominated shelf habitats contribute ~11 to 16 Tg C y-1. Since 2003, the Arctic seafloor area exposed to sunlight has increased by ~47,000 km2 y-1, expanding the realm of BPPs in a warming Arctic. Increased macrophyte abundance and productivity is expected along Arctic coastlines with continued ocean warming and sea ice loss. However, microalgal benthic primary production has increased in only a few shelf regions despite substantial sea ice loss over the past 20 y, as higher solar irradiance in the ice-free ocean is counterbalanced by reduced water transparency. This suggests complex impacts of climate change on Arctic light availability and marine primary production. Despite significant knowledge gaps on Arctic BPPs, their widespread presence and obvious contribution to coastal and shelf ecosystem production call for further investigation and for their inclusion in Arctic ecosystem models and carbon budgets.

Quantifying seaweed and seagrass beach deposits using high-resolution UAV imagery.

Macroalgae and seagrass wash ashore by tidal waters and episodic events and create an ocean-to-land transport of carbon and nutrients. On land, these deposits (beach wrack) are consumed by macrofauna, remineralized by microorganisms, or washed back to the sea, during which recycling of carbon and nitrogen affect the biochemical cycles in coastal zones. Manual quantification of beach wracks is time-consuming and often difficult due to complex topography and remote locations. Here, we present a novel method using Unoccupied Aerial Vehicle (UAV) photogrammetry combined with in situ measurements of carbon and nitrogen contents of wrack to quantify marine carbon and nutrient deposits in beach zones. The UAV method was tested against placed cubes ranging from 125 to 88,218 cm3 and demonstrated a high accuracy (R2 > 0.99) for volume acquisition when compared to manual measurements. Also, the UAV-based assessments of the cross-sectional area of beach deposits demonstrated a high accuracy when compared to manual and high-precision GNSS (Global Navigation Satellite System) measurements without significant differences between the methods. This demonstrated that UAVs can provide detailed spatial maps, three-dimensional (3D) surface models, and accurate volumetric assessments of beach wrack deposits. In three case studies, combined with carbon and nitrogen measures, total organic carbon and nitrogen deposits in beach wracks were quantified ranging from 4.3 to 9.7 and from 0.3 to 0.5 kg per meter coastline, respectively. In conclusion, this UAV method demonstrated an effective tool to quantify ecosystem carbon and nitrogen deposits relevant to ecosystem assessments and quantification of blue carbon stocks. The method is optimal when the terrain below beach wrack deposits is known, as in the case with before-and-after or repeated surveys. Further, UAVs display strong time- and cost-effective advantages over manual methods which is amplified with increasing project scale. We propose it as a valuable method for multiple scientific and commercial applications related to environmental monitoring and management, including marine resource exploration and exploitation.

Kelp carbon sink potential decreases with warming due to accelerating decomposition.

Cycling of organic carbon in the ocean has the potential to mitigate or exacerbate global climate change, but major questions remain about the environmental controls on organic carbon flux in the coastal zone. Here, we used a field experiment distributed across 28° of latitude, and the entire range of 2 dominant kelp species in the northern hemisphere, to measure decomposition rates of kelp detritus on the seafloor in relation to local environmental factors. Detritus decomposition in both species were strongly related to ocean temperature and initial carbon content, with higher rates of biomass loss at lower latitudes with warmer temperatures. Our experiment showed slow overall decomposition and turnover of kelp detritus and modeling of coastal residence times at our study sites revealed that a significant portion of this production can remain intact long enough to reach deep marine sinks. The results suggest that decomposition of these kelp species could accelerate with ocean warming and that low-latitude kelp forests could experience the greatest increase in remineralization with a 9% to 42% reduced potential for transport to long-term ocean sinks under short-term (RCP4.5) and long-term (RCP8.5) warming scenarios. However, slow decomposition at high latitudes, where kelp abundance is predicted to expand, indicates potential for increasing kelp-carbon sinks in cooler (northern) regions. Our findings reveal an important latitudinal gradient in coastal ecosystem function that provides an improved capacity to predict the implications of ocean warming on carbon cycling. Broad-scale patterns in organic carbon decomposition revealed here can be used to identify hotspots of carbon sequestration potential and resolve relationships between carbon cycling processes and ocean climate at a global scale.

Phytoplankton Productivity in an Arctic Fjord (West Greenland): Estimating Electron Requirements for Carbon Fixation and Oxygen Production.

Accurate quantification of pelagic primary production is essential for quantifying the marine carbon turnover and the energy supply to the food web. Knowing the electron requirement (Κ) for carbon (C) fixation (ΚC) and oxygen (O2) production (ΚO2), variable fluorescence has the potential to quantify primary production in microalgae, and hereby increasing spatial and temporal resolution of measurements compared to traditional methods. Here we quantify ΚC and ΚO2 through measures of Pulse Amplitude Modulated (PAM) fluorometry, C fixation and O2 production in an Arctic fjord (Godthåbsfjorden, W Greenland). Through short- (2h) and long-term (24h) experiments, rates of electron transfer (ETRPSII), C fixation and/or O2 production were quantified and compared. Absolute rates of ETR were derived by accounting for Photosystem II light absorption and spectral light composition. Two-hour incubations revealed a linear relationship between ETRPSII and gross 14C fixation (R2 = 0.81) during light-limited photosynthesis, giving a ΚC of 7.6 ± 0.6 (mean ± S.E.) mol é (mol C)-1. Diel net rates also demonstrated a linear relationship between ETRPSII and C fixation giving a ΚC of 11.2 ± 1.3 mol é (mol C)-1 (R2 = 0.86). For net O2 production the electron requirement was lower than for net C fixation giving 6.5 ± 0.9 mol é (mol O2)-1 (R2 = 0.94). This, however, still is an electron requirement 1.6 times higher than the theoretical minimum for O2 production [i.e. 4 mol é (mol O2)-1]. The discrepancy is explained by respiratory activity and non-photochemical electron requirements and the variability is discussed. In conclusion, the bio-optical method and derived electron requirement support conversion of ETR to units of C or O2, paving the road for improved spatial and temporal resolution of primary production estimates.

Aquatic eddy correlation: quantifying the artificial flux caused by stirring-sensitive O2 sensors.

In the last decade, the aquatic eddy correlation (EC) technique has proven to be a powerful approach for non-invasive measurements of oxygen fluxes across the sediment water interface. Fundamental to the EC approach is the correlation of turbulent velocity and oxygen concentration fluctuations measured with high frequencies in the same sampling volume. Oxygen concentrations are commonly measured with fast responding electrochemical microsensors. However, due to their own oxygen consumption, electrochemical microsensors are sensitive to changes of the diffusive boundary layer surrounding the probe and thus to changes in the ambient flow velocity. The so-called stirring sensitivity of microsensors constitutes an inherent correlation of flow velocity and oxygen sensing and thus an artificial flux which can confound the benthic flux determination. To assess the artificial flux we measured the correlation between the turbulent flow velocity and the signal of oxygen microsensors in a sealed annular flume without any oxygen sinks and sources. Experiments revealed significant correlations, even for sensors designed to have low stirring sensitivities of ~0.7%. The artificial fluxes depended on ambient flow conditions and, counter intuitively, increased at higher velocities because of the nonlinear contribution of turbulent velocity fluctuations. The measured artificial fluxes ranged from 2-70 mmol m(-2) d(-1) for weak and very strong turbulent flow, respectively. Further, the stirring sensitivity depended on the sensor orientation towards the flow. For a sensor orientation typically used in field studies, the artificial flux could be predicted using a simplified mathematical model. Optical microsensors (optodes) that should not exhibit a stirring sensitivity were tested in parallel and did not show any significant correlation between O2 signals and turbulent flow. In conclusion, EC data obtained with electrochemical sensors can be affected by artificial flux and we recommend using optical microsensors in future EC-studies.

Impacts of agricultural irrigation on nearby freshwater ecosystems: the seasonal influence of triazine herbicides in benthic algal communities.

A small hydrological basin (Lerma, NE Spain), transformed from its natural state (steppe) to rain-fed agriculture and recently to irrigation agriculture, has been monitored across four seasons of an agricultural year. The goal of this study was to assess how and whether agricultural activities impacted the nearby freshwater ecosystems via runoff. Specifically, we assessed the toxicity of three triazine herbicides, terbuthylazine, atrazine and simazine on the photosynthetic efficiency and structure of algal benthic biofilms (i.e., phototropic periphyton) in the small creek draining the basin. It was expected that the seasonal runoff of the herbicides in the creek affected the sensitivity of the periphyton in accord with the rationale of the Pollution Induced Community Tolerance (PICT): the exposure of the community to pollutants result in the replacement of sensitive species by more tolerant ones. In this way, PICT can serve to establish causal linkages between pollutants and the observed biological impacts. The periphyton presented significantly different sensitivities against terbuthylazine through the year in accord with the seasonal application of this herbicide in the crops nowadays. The sensitivity of already banned herbicides, atrazine and simazine does not display a clear seasonality. The different sensitivities to herbicides were in agreement with the expected exposures scenarios, according to the agricultural calendar, but not with the concentrations measured in water, which altogether indicates that the use of PICT approach may serve for long-term monitoring purposes. That will provide not only causal links between the occurrence of chemicals and their impacts on natural communities, but also information about the occurrence of chemicals that may escape from traditional sampling methods (water analysis). In addition, the EC50 and EC10 of periphyton for terbuthylazine or simazine are the first to be published and can be used for impact assessments.

System responses to equal doses of photosynthetically usable radiation of blue, green, and red light in the marine diatom Phaeodactylum tricornutum.

Due to the selective attenuation of solar light and the absorption properties of seawater and seawater constituents, free-floating photosynthetic organisms have to cope with rapid and unpredictable changes in both intensity and spectral quality. We have studied the transcriptional, metabolic and photo-physiological responses to light of different spectral quality in the marine diatom Phaeodactylum tricornutum through time-series studies of cultures exposed to equal doses of photosynthetically usable radiation of blue, green and red light. The experiments showed that short-term differences in gene expression and profiles are mainly light quality-dependent. Transcription of photosynthesis-associated nuclear genes was activated mainly through a light quality-independent mechanism likely to rely on chloroplast-to-nucleus signaling. In contrast, genes encoding proteins important for photoprotection and PSII repair were highly dependent on a blue light receptor-mediated signal. Changes in energy transfer efficiency by light-harvesting pigments were spectrally dependent; furthermore, a declining trend in photosynthetic efficiency was observed in red light. The combined results suggest that diatoms possess a light quality-dependent ability to activate photoprotection and efficient repair of photodamaged PSII. In spite of approximately equal numbers of PSII-absorbed quanta in blue, green and red light, the spectral quality of light is important for diatom responses to ambient light conditions.

Molecular and photosynthetic responses to prolonged darkness and subsequent acclimation to re-illumination in the diatom Phaeodactylum tricornutum.

Photosynthetic diatoms that live suspended throughout the water column will constantly be swept up and down by vertical mixing. When returned to the photic zone after experiencing longer periods in darkness, mechanisms exist that enable the diatoms both to survive sudden light exposure and immediately utilize the available energy in photosynthesis and growth. We have investigated both the response to prolonged darkness and the re-acclimation to moderate intensity white irradiance (E = 100 µmol m(-2) s(-1)) in the diatom Phaeodactylum tricornutum, using an integrated approach involving global transcriptional profiling, pigment analyses, imaging and photo-physiological measurements. The responses were studied during continuous white light, after 48 h of dark treatment and after 0.5 h, 6 h, and 24 h of re-exposure to the initial irradiance. The analyses resulted in several intriguing findings. Dark treatment of the cells led to 1) significantly decreased nuclear transcriptional activity, 2) distinct intracellular changes, 3) fixed ratios of the light-harvesting pigments despite a decrease in the total cell pigment pool, and 4) only a minor drop in photosynthetic efficiency (Φ(PSII_max)). Re-introduction of the cells to the initial light conditions revealed 5) distinct expression profiles for nuclear genes involved in photosynthesis and those involved in photoprotection, 6) rapid rise in photosynthetic parameters (α and rETR(max)) within 0.5 h of re-exposure to light despite a very modest de novo synthesis of photosynthetic compounds, and 7) increasingly efficient resonance energy transfer from fucoxanthin chlorophyll a/c-binding protein complexes to photosystem II reaction centers during the first 0.5 h, supporting the observations stated in 6). In summary, the results show that despite extensive transcriptional, metabolic and intracellular changes, the ability of cells to perform photosynthesis was kept intact during the length of the experiment. We conclude that P. tricornutum maintains a functional photosynthetic apparatus during dark periods that enables prompt recovery upon re-illumination.

An integrated analysis of molecular acclimation to high light in the marine diatom Phaeodactylum tricornutum.

Photosynthetic diatoms are exposed to rapid and unpredictable changes in irradiance and spectral quality, and must be able to acclimate their light harvesting systems to varying light conditions. Molecular mechanisms behind light acclimation in diatoms are largely unknown. We set out to investigate the mechanisms of high light acclimation in Phaeodactylum tricornutum using an integrated approach involving global transcriptional profiling, metabolite profiling and variable fluorescence technique. Algae cultures were acclimated to low light (LL), after which the cultures were transferred to high light (HL). Molecular, metabolic and physiological responses were studied at time points 0.5 h, 3 h, 6 h, 12 h, 24 h and 48 h after transfer to HL conditions. The integrated results indicate that the acclimation mechanisms in diatoms can be divided into an initial response phase (0-0.5 h), an intermediate acclimation phase (3-12 h) and a late acclimation phase (12-48 h). The initial phase is recognized by strong and rapid regulation of genes encoding proteins involved in photosynthesis, pigment metabolism and reactive oxygen species (ROS) scavenging systems. A significant increase in light protecting metabolites occur together with the induction of transcriptional processes involved in protection of cellular structures at this early phase. During the following phases, the metabolite profiling display a pronounced decrease in light harvesting pigments, whereas the variable fluorescence measurements show that the photosynthetic capacity increases strongly during the late acclimation phase. We show that P. tricornutum is capable of swift and efficient execution of photoprotective mechanisms, followed by changes in the composition of the photosynthetic machinery that enable the diatoms to utilize the excess energy available in HL. Central molecular players in light protection and acclimation to high irradiance have been identified.

TEMPERATURE EFFECTS ON MICROALGAL PHOTOSYNTHESIS-LIGHT RESPONSES MEASURED BY O2 PRODUCTION, PULSE-AMPLITUDE-MODULATED FLUORESCENCE, AND (14) C ASSIMILATION(1).

Short-term temperature effects on photosynthesis were investigated by measuring O2 production, PSII-fluorescence kinetics, and (14) C-incorporation rates in monocultures of the marine phytoplankton species Prorocentrum minimum (Pavill.) J. Schiller (Dinophyceae), Prymnesium parvum f. patelliferum (J. C. Green, D. J. Hibberd et Pienaar) A. Larsen (Coccolithophyceae), and Phaeodactylum tricornutum Bohlin (Bacillariophyceae), grown at 15°C and 80 µmol photons ·â€Šm(-2)  ·â€Šs(-1) . Photosynthesis versus irradiance curves were measured at seven temperatures (0°C-30°C) by all three approaches. The maximum photosynthetic rate (P(C) max ) was strongly stimulated by temperature, reached an optimum for Pro. minimum only (20°C-25°C), and showed a similar relative temperature response for the three applied methods, with Q10 ranging from 1.7 to 3.5. The maximum light utilization coefficient (α(C) ) was insensitive or decreased slightly with increasing temperature. Absolute rates of O2 production were calculated from pulse-amplitude-modulated (PAM) fluorometry measurements in combination with biooptical determination of absorbed quanta in PSII. The relationship between PAM-based O2 production and measured O2 production and (14) C assimilation showed a species-specific correlation, with 1.2-3.3 times higher absolute values of P(C) max and α(C) when calculated from PAM data for Pry. parvum and Ph. tricornutum but equivalent for Pro. minimum. The offset seemed to be temperature insensitive and could be explained by a lower quantum yield for O2 production than the theoretical maximum (due to Mehler-type reactions). Conclusively, the PAM technique can be used to study temperature responses of photosynthesis in microalgae when paying attention to the absorption properties in PSII.

RATE OF O2 PRODUCTION DERIVED FROM PULSE-AMPLITUDE-MODULATED FLUORESCENCE: TESTING THREE BIOOPTICAL APPROACHES AGAINST MEASURED O2 -PRODUCTION RATE(1).

Light absorption by phytoplankton is both species specific and affected by photoacclimational status. To estimate oxygenic photosynthesis from pulse-amplitude-modulated (PAM) fluorescence, the amount of quanta absorbed by PSII needs to be quantified. We present here three different biooptical approaches to estimate the fraction of light absorbed by PSII: (1) the factor 0.5, which implies that absorbed light is equally distributed among PSI and PSII; (2) the fraction of chl a in PSII, determined as the ratio between the scaled red-peak fluorescence excitation and the red absorption peak; and (3) the measure of light absorbed by PSII, determined from the scaling of the fluorescence excitation spectra to the absorption spectra by the "no-overshoot" procedure. Three marine phytoplankton species were used as test organisms: Prorocentrum minimum (Pavill.) J. Schiller (Dinophyceae), Prymnesium parvum cf. patelliferum (J. C. Green, D. J. Hibberd et Pienaar) A. Larsen (Haptophyceae), and Phaeodactylum tricornutum Bohlin (Bacillariophyceae). Photosynthesis versus irradiance (P vs. E) parameters calculated using the three approaches were compared with P versus E parameters obtained from simultaneously measured rates of oxygen production. Generally, approach 1 underestimated, while approach 2 overestimated the gross O2 -production rate calculated from PAM fluorescence. Approach 3, in principle the best approach to estimate quanta absorbed by PSII, was also superior according to observations. Hence, we recommend approach 3 for estimation of gross O2 -production rates based on PAM fluorescence measurements.