Stefan G. H. Simis

Estimating phytoplankton primary production in Lake IJsselmeer (The Netherlands) using variable fluorescence (PAM-FRRF) and C-uptake techniques

In this paper we compare measured rates of C-fixation obtained using in situ deployment of a fast repetition rate fluorometer (FRRF) applying two different calculation protocols and a waterPAM fluorometer. The measurements were carried out in the large, shallow Lake IJsselmeer (The Netherlands). Turbidity appears to be the most important factor in explaining photosynthetic parameters. Non-photochemical quenching (NPQ) in the antenna was observed, and the degree of NPQ was positively correlated with the irradiance and mixing conditions. However, NPQ did not affect the functional cross-section. Conversion of photosynthetic electron transport rates (ETR) into rates of carbon fixation requires knowledge of two or three parameters (depending on the protocol and instrument used) to calculate C-fixation from ETR: the electron yield (Φe, i.e. the reciprocal of the quantum requirement of PSII), the photosynthetic quotient (PQ) and the number of PSII (nPSII). Using normally assumed values for these three parameters resulted in overestimation of the measured rate of C-fixation but, in all cases, ETR was linearly related to it. The degree of overestimation was rather constant, despite considerable changes in phytoplankton composition. Spectral correction of fluorescence data caused a reduction of 30% or an increase of 44% in the estimated C-fixation, depending how C-fixation was estimated. We describe a method that allows estimation of the Φe/PQ ratio based on a comparison of the quantum efficiencies for C-fixation and PSII and show that, with a single and realistic value for this ratio, primary production could be accurately predicted at different times of the year.

Predicting the Electron Requirement for Carbon Fixation in Seas and Oceans

Marine phytoplankton account for about 50% of all global net primary productivity (NPP). Active fluorometry, mainly Fast Repetition Rate fluorometry (FRRf), has been advocated as means of providing high resolution estimates of NPP. However, not measuring CO2-fixation directly, FRRf instead provides photosynthetic quantum efficiency estimates from which electron transfer rates (ETR) and ultimately CO2-fixation rates can be derived. Consequently, conversions of ETRs to CO2-fixation requires knowledge of the electron requirement for carbon fixation (Φe,C, ETR/CO2 uptake rate) and its dependence on environmental gradients. Such knowledge is critical for large scale implementation of active fluorescence to better characterise CO2-uptake. Here we examine the variability of experimentally determined Φe,C values in relation to key environmental variables with the aim of developing new working algorithms for the calculation of Φe,C from environmental variables. Coincident FRRf and 14C-uptake and environmental data from 14 studies covering 12 marine regions were analysed via a meta-analytical, non-parametric, multivariate approach. Combining all studies, Φe,C varied between 1.15 and 54.2 mol e− (mol C)−1 with a mean of 10.9±6.91 mol e− mol C)−1. Although variability of Φe,C was related to environmental gradients at global scales, region-specific analyses provided far improved predictive capability. However, use of regional Φ e,C algorithms requires objective means of defining regions of interest, which remains challenging. Considering individual studies and specific small-scale regions, temperature, nutrient and light availability were correlated with Φ e,C albeit to varying degrees and depending on the study/region and the composition of the extant phytoplankton community. At the level of large biogeographic regions and distinct water masses, Φ e,C was related to nutrient availability, chlorophyll, as well as temperature and/or salinity in most regions, while light availability was also important in Baltic Sea and shelf waters. The novel Φ e,C algorithms provide a major step forward for widespread fluorometry-based NPP estimates and highlight the need for further studying the natural variability of Φe,C to verify and develop algorithms with improved accuracy.

Implementing and Innovating Marine Monitoring Approaches for Assessing Marine Environmental Status

Marine environmental monitoring has tended to focus on site-specific methods of investigation. These traditional methods have low spatial and temporal resolution and are relatively labor intensive per unit area/time that they cover. To implement the Marine Strategy Framework Directive (MSFD), European Member States are required to improve marine monitoring and design monitoring networks. This can be achieved by developing and testing innovative and cost-effective monitoring systems, as well as indicators of environmental status. Here, we present several recently developed methodologies and technologies to improve marine biodiversity indicators and monitoring methods. The innovative tools are discussed concerning the technologies presently utilized as well as the advantages and disadvantages of their use in routine monitoring. In particular, the present analysis focuses on: (i) molecular approaches, including microarray, Real Time quantitative PCR (qPCR), and metagenetic (metabarcoding) tools; (ii) optical (remote) sensing and acoustic methods; and (iii) in situ monitoring instruments. We also discuss their applications in marine monitoring within the MSFD through the analysis of case studies in order to evaluate their potential utilization in future routine marine monitoring. We show that these recently-developed technologies can present clear advantages in accuracy, efficiency and cost.

Interaction Effects of Light, Temperature and Nutrient Limitations (N, P and Si) on Growth, Stoichiometry and Photosynthetic Parameters of the Cold-Water Diatom Chaetoceros wighamii

Light (20-450 μmol photons m-2 s-1), temperature (3-11°C) and inorganic nutrient composition (nutrient replete and N, P and Si limitation) were manipulated to study their combined influence on growth, stoichiometry (C:N:P:Chl a) and primary production of the cold water diatom Chaetoceros wighamii. During exponential growth, the maximum growth rate (~0.8 d-1) was observed at high temperture and light; at 3°C the growth rate was ~30% lower under similar light conditions. The interaction effect of light and temperature were clearly visible from growth and cellular stoichiometry. The average C:N:P molar ratio was 80:13:1 during exponential growth, but the range, due to different light acclimation, was widest at the lowest temperature, reaching very low C:P (~50) and N:P ratios (~8) at low light and temperature. The C:Chl a ratio had also a wider range at the lowest temperature during exponential growth, ranging 16-48 (weight ratio) at 3°C compared with 17-33 at 11°C. During exponential growth, there was no clear trend in the Chl a normalized, initial slope (α*) of the photosynthesis-irradiance (PE) curve, but the maximum photosynthetic production (Pm) was highest for cultures acclimated to the highest light and temperature. During the stationary growth phase, the stoichiometric relationship depended on the limiting nutrient, but with generally increasing C:N:P ratio. The average photosynthetic quotient (PQ) during exponential growth was 1.26 but decreased to <1 under nutrient and light limitation, probably due to photorespiration. The results clearly demonstrate that there are interaction effects between light, temperature and nutrient limitation, and the data suggests greater variability of key parameters at low temperature. Understanding these dynamics will be important for improving models of aquatic primary production and biogeochemical cycles in a warming climate.

Dynamic modelling of viral impact on cyanobacterial populations in shallow lakes: Implications of burst size

Laboratory experiments with whole water-columns from shallow, eutrophic lakes repeatedly showed collapse of the predominant filamentous cyanobacteria. The collapse could be due to viral activity, from the evidence of electron microscopy of infected cyanobacterial cells and observed dynamics of virus-like particles. Burst-size effects on single-host single-virus dynamics was modelled for nutrient-replete growth of the cyanobacteria and fixed viral decay rate in the water column. The model combined previously published equations for nutrient-replete cyanobacterial growth and virus–host relationship. According to the model results, burst sizes greater than 200 to 400 virions per cell would result in host extinction, whereas lower numbers would allow coexistence, and even stable population densities of host and virus. High-nutrient status of the host cells might accommodate a large burst size. The ecological implication could be that burst-size increase accompanying a transition from phosphorus to light-limited cyanobacterial growth might destabilize the virus–host interaction and result in the population collapse observed in the experiments.

Validation of MERIS Case-2 Water Products in Lake Taihu, China

Three artificial neural network (ANN) processors available as plug-in modules for the Basic ERS & ENVISAT (A)ATSR and MERIS Toolbox (BEAM) were validated in Lake Taihu, China. Mean deviations of reflectance derived from Lakes\Boreal and Lakes\Eutrophic were 10-90%, while reflectance from the FUB-WeW processor showed larger errors. All processors showed underestimates of chlorophyll a (Chl-a), total suspended matter (TSM), and phytoplankton pigment absorption, while particulate scattering values were severely overestimated. None of the readily available MERIS processors is currently able to separate atmospheric and water-leaving radiance over Lake Taihu, while the retrieval of phytoplankton biomass through ANN processors shows promise.

Contrasting seasonality in optical-biogeochemical properties of the Baltic Sea

Optical-biogeochemical relationships of particulate and dissolved organic matter are presented in support of remote sensing of the Baltic Sea pelagic. This system exhibits strong seasonality in phytoplankton community composition and wide gradients of chromophoric dissolved organic matter (CDOM), properties which are poorly handled by existing remote sensing algorithms. Absorption and scattering properties of particulate matter reflected the seasonality in biological (phytoplankton succession) and physical (thermal stratification) processes. Inherent optical properties showed much wider variability when normalized to the chlorophyll-a concentration compared to normalization to either total suspended matter dry weight or particulate organic carbon. The particle population had the largest optical variability in summer and was dominated by organic matter in both seasons. The geographic variability of CDOM and relationships with dissolved organic carbon (DOC) are also presented. CDOM dominated light absorption at blue wavelengths, contributing 81% (median) of the absorption by all water constituents at 400 nm and 63% at 442 nm. Consequentially, 90% of water-leaving radiance at 412 nm originated from a layer (z90) no deeper than approximately 1.0 m. With water increasingly attenuating light at longer wavelengths, a green peak in light penetration and reflectance is always present in these waters, with z90 up to 3.0–3.5 m depth, whereas z90 only exceeds 5 m at biomass < 5 mg Chla m-3. High absorption combined with a weakly scattering particle population (despite median phytoplankton biomass of 14.1 and 4.3 mg Chla m-3 in spring and summer samples, respectively), characterize this sea as a dark water body for which dedicated or exceptionally robust remote sensing techniques are required. Seasonal and regional optical-biogeochemical models, data distributions, and an extensive set of simulated remote-sensing reflectance spectra for testing of remote sensing algorithms are provided as supplementary data.

Atmospheric Correction Performance of Hyperspectral Airborne Imagery over a Small Eutrophic Lake under Changing Cloud Cover

Atmospheric correction of remotely sensed imagery of inland water bodies is essential to interpret water-leaving radiance signals and for the accurate retrieval of water quality variables. Atmospheric correction is particularly challenging over inhomogeneous water bodies surrounded by comparatively bright land surface. We present results of AisaFENIX airborne hyperspectral imagery collected over a small inland water body under changing cloud cover, presenting challenging but common conditions for atmospheric correction. This is the first evaluation of the performance of the FENIX sensor over water bodies. ATCOR4, which is not specifically designed for atmospheric correction over water and does not make any assumptions on water type, was used to obtain atmospherically corrected reflectance values, which were compared to in situ water-leaving reflectance collected at six stations. Three different atmospheric correction strategies in ATCOR4 was tested. The strategy using fully image-derived and spatially varying atmospheric parameters produced a reflectance accuracy of ±0.002, i.e., a difference of less than 15% compared to the in situ reference reflectance. Amplitude and shape of the remotely sensed reflectance spectra were in general accordance with the in situ data. The spectral angle was better than 4.1° for the best cases, in the spectral range of 450–750 nm. The retrieval of chlorophyll-a (Chl-a) concentration using a popular semi-analytical band ratio algorithm for turbid inland waters gave an accuracy of ~16% or 4.4 mg/m3 compared to retrieval of Chl-a from reflectance measured in situ. Using fixed ATCOR4 processing parameters for whole images improved Chl-a retrieval results from ~6 mg/m3 difference to reference to approximately 2 mg/m3. We conclude that the AisaFENIX sensor, in combination with ATCOR4 in image-driven parametrization, can be successfully used for inland water quality observations. This implies that the need for in situ reference measurements is not as strict as has been assumed and a high degree of automation in processing is possible.

Validation of a spectral correction procedure for sun and sky reflections in above-water reflectance measurements

A three-component reflectance model (3C) is applied to above-water radiometric measurements to derive remote-sensing reflectance Rrs (λ). 3C provides a spectrally resolved offset Δ(λ) to correct for residual sun and sky radiance (Rayleigh- and aerosol-scattered) reflections on the water surface that were not represented by sky radiance measurements. 3C is validated with a data set of matching above- and below-water radiometric measurements collected in the Baltic Sea, and compared against a scalar offset correction Δ. Correction with Δ(λ) instead of Δ consistently reduced the (mean normalized root-mean-square) deviation between Rrs (λ) and reference reflectances to comparable levels for clear (Δ: 14.3 ± 2.5 %, Δ(λ): 8.2 ± 1.7 %), partly clouded (Δ: 15.4 ± 2.1 %, Δ(λ): 6.5 ± 1.4 %), and completely overcast (Δ: 10.8 ± 1.7 %, Δ(λ): 6.3 ± 1.8 %) sky conditions. The improvement was most pronounced under inhomogeneous sky conditions when measurements of sky radiance tend to be less representative of surface-reflected radiance. Accounting for both sun glint and sky reflections also relaxes constraints on measurement geometry, which was demonstrated based on a semi-continuous daytime data set recorded in a eutrophic freshwater lake in the Netherlands. Rrs (λ) that were derived throughout the day varied spectrally by less than 2 % relative standard deviation. Implications on measurement protocols are discussed. An open source software library for processing reflectance measurements was developed and is made publicly available.

Variability of adjacency effects in sky reflectance measurements

Sky reflectance Rsky(λ) is used to correct in situ reflectance measurements in the remote detection of water color. We analyzed the directional and spectral variability in Rsky(λ) due to adjacency effects against an atmospheric radiance model. The analysis is based on one year of semi-continuous Rsky(λ) observations that were recorded in two azimuth directions. Adjacency effects contributed to Rsky(λ) dependence on season and viewing angle and predominantly in the near-infrared (NIR). For our test area, adjacency effects spectrally resembled a generic vegetation spectrum. The adjacency effect was weakly dependent on the magnitude of Rayleigh- and aerosol-scattered radiance. The reflectance differed between viewing directions 5.4±6.3% for adjacency effects and 21.0±19.8% for Rayleigh- and aerosol-scattered Rsky(λ) in the NIR. Under which conditions in situ water reflectance observations require dedicated correction for adjacency effects is discussed. We provide an open source implementation of our method to aid identification of such conditions.