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Existing statistical models of in vivo light absorption by phytoplankton (Woźniak & Ostrowska 1990, Bricaud et al. 1995, 1998) describe the dependence of the phytoplankton specific spectral absorption coefficient a∗ pl(λ) on the chlorophyll a concentration Ca in seawater. However, the models do not take into account the variability in this relationship due to phytoplankton acclimation. The observed variability in the light absorption coefficient and its components due to various pigments with depth and geographical position at sea, requires further accurate modelling in order to improve satellite remote sensing algorithms and interpretation of ocean colour maps. The aim of this paper is to formulate an improved model of the phytoplankton spectral absorption capacity which takes account of the pigment composition and absorption changes resulting from photo- and chromatic acclimation processes, and the pigment package effect. It is a synthesis of earlier models and the following statistical generalisations: (1) statistical relationships between various pigment group concentrations and light field properties in the sea (described by Majchrowski & Ostrowska 2000, this volume); (2) a model of light absorption by phytoplankton capable of determining the mathematical relationships between the spectral absorption coefficients of the various photosynthetic and photoprotecting pigment groups, and their concentrations in seawater (Woźniak et al. 1999); (3) bio-optical models of light propagation in oceanic Case 1 Waters and Baltic Case 2 Waters (Woźniak et al. 1992a, b, 1995a,b). The generalised model described in this paper permits the total phytoplankton light absorption coefficient in vivo as well as its components related to the various photosynthetic and photoprotecting pigments to be determined using only the surface irradiance PAR(0+) surface chlorophyll concentration Ca(0) and depth z in the sea as input data.
The aim of this work was to find statistical relationships between the concentrations of accessory pigments in natural populations of marine phytoplankton and the absolute levels and spectral distributions of underwater irradiance. To this end, empirical data sets from some 600 stations in different parts of the seas and oceans were analysed. These data were obtained from the authors’ own research and from the Internet’s bio-optical data base. They included the vertical distributions of the concentrations of various pigments (identified chromatographically) and the vertical and spectral distributions of the underwater irradiance measured in situ or determined indirectly from bio-optical models. The analysis covered a total of some 4000 points illustrating the dependence of pigment concentration on underwater irradiance characteristics, corresponding to different depths in the sea. The analysis showed that the factor governing the occurrence of photoprotecting carotenoids (PPC) is short-wave radiation λ < 480 nm. A mathematical relationship was established between the relative PPC concentration (relative with respect to the chlorophyll a concentration) and the magnitude of the absorbed radiative energy per unit mass of chlorophyll a from the spectral interval λ < 480 nm, averaged in the water layers Δz = 60 m (or less near the surface) to account for vertical mixing. This absorbed short-wave radiation (λ < 480 nm) was given the name of Potentially Destructive Radiation (PDR∗(z)). Analysis of the relationships between the concentrations of particular photosynthetic pigments (PSP), i.e. chlorophyll b, chlorophyll c, photosynthetic carotenoids (PSC), and the underwater irradiance characteristics indicated that these concentrations were only slightly dependent on the absolute level of irradiance E0(λ), but that they depended strongly on the relative spectral distribution of this irradiance f(λ) = E0(λ)/PAR0. The relevant approximate statistical relationships between the relative concentrations of particular PSP and the function of spectral fitting Fj , averaged in the layer Δz, were derived. Certain statistical relationships between the pigment composition of the phtyoplankton and the irradiance field characteristics are due to the photo- and chromatic acclimation of natural populations of marine phytoplankton. These relationships can be applied in models of the coefficients of light absorption by phytoplankton.
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The overriding and far-reaching aim of our work has been to achieve a good understanding of the processes of light interaction with phytoplankton in the sea and to develop an innovative physical model of photosynthesis in the marine environment,suita ble for the remote sensing of marine primary production. Unlike previous models,the present one takes greater account of the complexity of the physiological processes in phytoplankton. We have focused in particular on photophysiological processes,whic h are governed directly or indirectly by light energy,or in which light, besides the nutrient content in and the temperature of seawater,is one of the principal limiting factors. To achieve this aim we have carried out comprehensive statistical analyses of the natural variability of the main photophysiological properties of phytoplankton and their links with the principal abiotic factors in the sea. These analyses have made use of extensive empirical data gathered in a wide diversity of seas and oceans by Polish and Russian teams as well as by joint Polish-Russian expeditions. Data sets available on the Internet have also been applied. As a result,a set of more or less complex,semi-empir ical models of light-stimulated processes occurring in marine phytoplankton cells has been developed. The trophic type of sea, photo-acclimation and the production of photoprotecting carotenoids,c hromatic acclimation and the production of various forms of chlorophyll-antennas and photosynthetic carotenoids,cell adaptation by the package effect, light absorption, photosynthesis, photoinhibition,the fluorescence effect,a nd the activation of PS2 centres are all considered in the models. These take into account not only the influence of light,but also, indirectly,tha t of the vertical mixing of water; in the case of photosynthesis,the quantum yield has been also formulated as being dependent on the nutrient concentrations and the temperature of seawater. The bio-optical spectral models of irradiance transmittance in case 1 oceanic waters and case 2 Baltic waters,dev eloped earlier,a lso are described in this paper. The development of the models presented here is not yet complete and they all need continual improvement. Nevertheless,w e have used them on a preliminary basis for calculating various photosynthetic characteristics at different depths in the sea,su ch as the concentration of chlorophyll and other pigments, and primary production. The practical algorithm we have constructed allows the vertical distribution of these characteristics to be determined from three input data: chlorophyll a concentration,irradiance, and temperature at the sea surface. Since all three data can be measured remotely,ou r algorithm can be applied as the ‘marine part’ of the remote sensing algorithms used for detecting marine photosynthesis.
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