Oceanologia No. 42 (2) / 00

Contents

Preface

Papers

Papers


Luminescence and photosynthesis of marine phytoplankton - a brief presentation of new results
Oceanologia 2000, 42 (2), 137-156

Bogdan Woźniak1, 2, Jerzy Dera1
1Institute of Oceanology, Polish Academy of Sciences, Powstańców Warszawy 55, PL-81-712 Sopot, Poland; wozniak@iopan.gda.pl
2Institute of Physics, Pedagogical University, Arciszewskiego 22 B, PL-76-200 Słupsk, Poland;

Keywords: phytoplankton, photoprotecting pigments, photosynthetic pigments, phytoplankton light absorption, phytoplankton fluorescence, photosynthesis quantum yield

Manuscript received 13 March 2000, reviewed 30 March 2000, accepted 10 April 2000.
Abstract
This volume contains a set of eight papers presenting the results of the latest research into the interaction of light with marine phytoplankton by teams from the Marine Physics Department at the IO PAS in Sopot, and the Department of Environmental Physics at the Pedagogical University of Słupsk. These results were presented at the 'Second Workshop on Luminescence and Photosynthesis of Marine Phytoplankton' (Sopot-Paraszyno, 11-15 October 1999) sponsored by the Polish State Committee for Scientific Research. This introductory article discusses the most important assumptions and objectives of the research, and outlines the latest results. These are subsequently discussed in detail in the following papers: (1) Majchrowski & Ostrowska, Influence of photo- and chromatic acclimation on pigment composition in the sea, (2) Woźniak et al., Model of the 'in vivo' spectral absorption of algal pigments. Part 1. Mathematical apparatus, (3) Majchrowski et al., Model of the 'in vivo'spectral absorption of algal pigments. Part 2. Practical applications of the model, (4) Ostrowska et al., Variability of the specific fluorescence of chlorophyll in the ocean. Part 1. Theory of classical 'in situ' chlorophyll fluorometry, (5) Ostrowska et al., Variability of the specific fluorescence of chlorophyll in the ocean. Part 2. Fluorometric method of chlorophyll a determination, (6) Ficek et al., Influence of non-photosynthetic pigments on the measured quantum yield of photosynthesis, (7) Ficek et al., Variability of the portion of functional PS2 reaction centres in the light of a fluorometric study.
For the reader's convenience, we append a list of the symbols denoting the physical quantities used in the texts. The nomenclature and denotations are in line with the conventions employed in the subject literature.
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Influence of photo- and chromatic acclimation on pigment composition in the sea
Oceanologia 2000, 42 (2), 157-175

Roman Majchrowski2, Mirosława Ostrowska1
1Institute of Oceanology, Polish Academy of Sciences, Powstańców Warszawy 55, PL-81-712 Sopot, Poland;
2Institute of Physics, Pedagogical University, Arciszewskiego 22 B, PL-76-200 Słupsk, Poland; roman@wsp.slupsk.pl

Keywords: phytoplankton pigments, pigments light absorption, photoprotecting pigments, photosynthetic pigments, photoacclimation, chromatic acclimation

Manuscript received 9 March 2000, reviewed 27 March 2000, accepted 7 April 2000.
Abstract
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 c z = 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 phytoplankton 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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Model of the in vivo spectral absorption of algal pigments. Part 1. Mathematical apparatus
Oceanologia 2000, 42 (2), 177-190

Bogdan Woźniak1, 2, Jerzy Dera1, Dariusz Ficek2, Roman Majchrowski2, Sławomir Kaczmarek1, Mirosława Ostrowska1, Olga I. Koblentz-Mishke3
1Institute of Oceanology, Polish Academy of Sciences, Powstańców Warszawy 55, PL-81-712 Sopot, Poland; wozniak@iopan.gda.pl
2Institute of Physics, Pedagogical University, Arciszewskiego 22 B, PL-76-200 Słupsk, Poland;
3P. P. Shirshov Institute of Oceanology, Russian Academy of Sciences, Krasikova 23, Moscow, 117218 Russia;

Keywords: phytoplankton light absorption, photoprotecting pigments, photosynthetic pigments, photoacclimation, chromatic acclimation, bio-optical modelling

Manuscript received 15 March 2000, reviewed 5 April 2000, accepted 17 April 2000.
Abstract
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.
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Model of the in vivo spectral absorption of algal pigments. Part 2. Practical applications of the model
Oceanologia 2000, 42 (2), 191-202

Roman Majchrowski2, Bogdan Woźniak1, 2, Jerzy Dera1, Dariusz Ficek2, Sławomir Kaczmarek1, Mirosława Ostrowska1, Olga I. Koblentz-Mishke3
1Institute of Oceanology, Polish Academy of Sciences, Powstańców Warszawy 55, PL-81-712 Sopot, Poland;
2Institute of Physics, Pedagogical University, Arciszewskiego 22 B, PL-76-200 Słupsk, Poland; roman@wsp.slupsk.pl
3P. P. Shirshov Institute of Oceanology, Russian Academy of Sciences, Krasikova 23, Moscow, 117218 Russia;

Keywords: phytoplankton, light absorption, bio-optical modelling

Manuscript received 2 March 2000, reviewed 20 March 2000, accepted 27 March 2000.
Abstract
The article describes applications and accuracy analyses of a statistical model of light absorption by phytoplankton that accounts for the influence of photo- and chromatic acclimation on its absorption properties. Part 1 of this work (see Woźniak et al. 2000, this volume) describes the mathematical apparatus of the model. Earlier models by Woźniak & Ostrowska (1990) and by Bricaud et al. (1995, 1998) are analysed for comparison. Empirical verification of these three models shows that the new model provides a much better approximation of phytoplankton absorption properties than do the earlier models. The statistical errors in estimating the mean absorption coefficient apl, for example, are σ+ = 36% for the new model, whereas for the earlier models the figures are σ+ = 43% (Bricaud et al. 1995, 1998) and σ+ = 59% (Woźniak & Ostrowska 1990). Example applications are given of the new model illustrating the variability in phytoplankton absorption properties with depth and trophicity of the sea.
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Variability of the specific fluorescence of chlorophyll in the ocean. Part 1. Theory of classical in situ chlorophyll fluorometry
Oceanologia 2000, 42 (2), 203-219

Mirosława Ostrowska1, Roman Majchrowski2, Dimitrii N. Matorin3, Bogdan Woźniak1, 2
1Institute of Oceanology, Polish Academy of Sciences, Powstańców Warszawy 55, PL-81-712 Sopot, Poland; ostra@iopan.gda.pl
2Institute of Physics, Pedagogical University, Arciszewskiego 22 B, PL-76-200 Słupsk, Poland;
3Department of Biophysics, Faculty of Biology, Moscow State University, Moscow, 117218 Russia;

Keywords: plant luminescence, phytoplankton fluorescence in the ocean, specific chlorophyll fluorescence in vivo, theory of classical fluorometry, fluorometric method

Manuscript received 17 March 2000, reviewed 12 April 2000, accepted 30 April 2000.
Abstract
The range of variability of the fluorescence properties of marine phytoplankton in different trophic types of seas and at different depths in the sea is analysed theoretically. An attempt is also made to interpret artificially induced in situ fluorescence measured with submersible fluorometers. To do this, earlier optical models of light absorption by phytoplankton (see Woźniak et al. 2000, this volume) and actual empirical data were applied. A straightforward theoretical model of artificially photoinduced phytoplankton fluorescence accounting for the complex influence of different photophysiological characteristics of phytoplankton and the optical characteristics of the instrument has been worked out. A physical method of determining chlorophyll a concentrations in seawater from fluorescence measured in situ with contact fluorometers can be based on this model.
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Variability of the specific fluorescence of chlorophyll in the ocean. Part 2. Fluorometric method of chlorophyll a determination
Oceanologia 2000, 42 (2), 221-229

Mirosława Ostrowska
Institute of Oceanology, Polish Academy of Sciences, Powstańców Warszawy 55, PL-81-712 Sopot, Poland; ostra@iopan.gda.pl
Dimitrii N. Matorin
Department of Biophysics, Faculty of Biology, Moscow State University, Moscow, 117218 Russia;
Dariusz Ficek
Institute of Physics, Pedagogical University, Arciszewskiego 22 B, PL-76-200 Słupsk, Poland;


Keywords: plant luminescence, chlorophyll adetermination, fluorometric method

Manuscript received 22 March 2000, reviewed 18 April 2000, accepted 4 May 2000.
Abstract
Two methods of determining the chlorophyll a concentration in the sea have been formulated on the basis of artificially induced fluorescence measured with the aid of submersible fluorometers. The method of statistical correlation is founded on the empirical relationship between fluorescence and chlorophyll concentration. The theoretical model of fluorescence described in Part 1 of this paper (see Ostrowska et al. 2000, this volume) provides the basis of the other method, the physical method. This describes the dependence of the specific fluorescence of phytoplankton on the chlorophyll concentration, a diversity of photophysiological properties of phytoplankton and the optical characteristics of the fluorometer. In order to assess their practicability, the methods were subjected to empirical verification. This showed that the physical method yielded chlorophyll concentrations of far greater accuracy. The respective error factors of the estimated chlorophyll concentration were x = 2.07 for the correlation method and x = 1.5 for the physical method. This means that the statistical logarithmic error varies from -52 to +107% in the case of the former method but only from -33 to +51% in the case of the latter. Thus, modifying the methodology has much improved the accuracy of chlorophyll determinations.
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Influence of non-photosynthetic pigments on the measured quantum yield of photosynthesis
Oceanologia 2000, 42 (2), 231-242

Dariusz Ficek2, Roman Majchrowski2, Mirosława Ostrowska1, Bogdan Woźniak1, 2
1Institute of Oceanology, Polish Academy of Sciences, Powstańców Warszawy 55, PL-81-712 Sopot, Poland;
2Institute of Physics, Pedagogical University, Arciszewskiego 22 B, PL-76-200 Słupsk, Poland; darek@wsp.slupsk.pl

Keywords: photosynthesis quantum yield, non-photosynthetic pigment factor, bio-optical model

Manuscript received 20 March 2000, reviewed 10 April 2000, accepted 19 April 2000.
Abstract
The aim of this work was to assess the effect of non-photosynthetic (photoprotecting) pigments on the measured quantum yield of photosynthesis in the sea. The energy absorbed by these pigments is not utilised during photosynthesis. As a result, the measured yield of this process, i.e. the photosynthetic yield referred to the total energy absorbed by all phytoplankton pigments, is less than the actual quantum yield of photosynthesis, i.e. the yield referred to the energy absorbed by photosynthetic pigments only. The model of the absorption properties of marine phytoplankton derived by the authors (see Woźniak et al. 2000, this volume) was employed to determine the relevant contributions of photosynthetic and non-photosynthetic pigments to the total energy absorbed by phytoplankton in different trophic types of seas and at different depths in the water column. On this basis the non-photosynthetic pigment absorption factor fa, which describes the relation between the true and measured quantum yields of photosynthesis, could be characterised. The analysis shows that fa varies in value from 0.33 to 1, and that it depends on the trophic type of sea and the depth in the water column. The values of this factor are usually highest in eutrophic waters and decrease as waters become progressively more oligotrophic. It is also characteristic of fa that it increases with increasing depth in the sea.
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Variability of the portion of functional PS2 reaction centres in the light of a fluorometric study
Oceanologia 2000, 42 (2), 243-250

Dariusz Ficek2, Mirosława Ostrowska1, Malgorzata Kuzio2, Sergey I. Pogosyan3
1Institute of Oceanology, Polish Academy of Sciences, Powstańców Warszawy 55, PL-81-712 Sopot, Poland;
2Institute of Physics, Pedagogical University, Arciszewskiego 22 B, PL-76-200 Słupsk, Poland; e-mail: darek@wsp.slupsk.pl
3Department of Biophysics, Faculty of Biology, Moscow State University, Moscow, 117218 Russia;

Keywords: quantum yield of photosynthesis, portion of functional PS2 reaction centres, fluorometric method

Manuscript received 6 March 2000, reviewed 24 March 2000, accepted 31 March 2000.
Abstract
The paper contains a preliminary analysis of the links between the portion fc of functional PS2 reaction centres in the photosynthetic apparatus of marine phytoplankton and environmental factors. The analysis is based inter alia on fluorometric measurements of fc (see Kolber & Falkowski 1993) in water sampled from different depths and trophic types of sea. From the statistical generalisations was derived an analytical form of the relationship between fc, and the optical depth and trophic type of sea (the trophicity index was taken to be the surface concentration of chlorophyll a). According to this relationship, fc rises as the trophicity of the sea does so. Moreover, there is a certain optimal optical depth for each type of water at which the number of functional PS2 reaction centres reaches a maximum. Above or below this depth the value of fc falls. At the present stage of investigations it seems reasonable to assume that this drop in the number of functional PS2 reaction centres close to the surface is due to the destructive effect of excessive irradiance. On the other hand, their reduced number at greater depths, below the fc maximum, can be attributed to the deficit of light and the consequent destruction of reaction centres.
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Sky glint correction in measurements of upward radiance above the sea surface
Oceanologia 2000, 42 (2), 251-262

Jerzy Olszewski, Piotr Kowalczuk
Institute of Oceanology, Polish Academy of Sciences, Powstańców Warszawy 55, PL-81-712 Sopot, Poland; e-mail: olszewsk@iopan.gda.pl

Keywords: marine optics, remote sensing, glint reduction

Manuscript received 10 February 2000, reviewed 7 April 2000, accepted 17 April 2000.
Abstract
An experiment has been performed to determine the upward water-leaving radiance by non-contact measurement of the total upward and downward radiance above the sea surface from a moving ship. The method for achieving this aim is described: the radiance meters are both tilted in such a way that the upward radiance meter can 'see' that part of the measured downward radiance which would be reflected if the water surface were smooth and which is not derived directly from solar glitter. Both meters are firmly fixed in a special frame, which ensures that the required orientation is the most probable one. Time records of the measured parameters are analysed. The results are presented in several forms: frequency (histogram) analysis appears to be the most promising one.
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In situ and simulated in situ primary production in the Gulf of Gdańsk
Oceanologia 2000, 42 (2), 263-282

Henryk Renk, Stanisław Ochocki, Sławomir Kurzyk
Sea Fisheries Institute, H. Kollataja 1, PL-81-332, Gdynia, Poland; e-mail: sochocki@mir.gdynia.pl

Keywords: primary production, chlorophyll a, assimilation number, light curves of phytoplankton, Baltic Sea

Manuscript received 11 Febuary 2000, reviewed 15 May 2000, accepted 19 May 2000.
Abstract
The method discussed in this article allows the simulated in situ primary production in the southern Baltic to be evaluated. To estimate the daily primary production at a~given field station, the following parameters have to be measured: the coefficients AN and Es (constants from the photosynthesis-light curve for phytoplankton), and the scalar irradiance attenuation coefficient (k), chlorophyll a concentration (Chl) and daily irradiation just below the sea surface (PAR). The results of simulated in situ primary production are in good agreement with the in situ measurements.
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