Seasonal variation in light- and temperature-dependent growth of marine planktonic diatoms in in situ dialysis cultures in the Trondheimsfjord,norway (63°N)

Three species of diatoms, Skeletonema costatum (Grev.) Cleve, Thalassiosira gravida Cleve, and T. pseudonana (Hustedt) Hasle et Heimdal, were grown in in situ dialysis culture in the Trondheimsfjord at depths of 0.5 and 4 m. The rates of growth and the chemical composition of exponentially growing cells were monitored and related to seasonal changes in illumination and temperature. Functions correlating growth rate with temperature were deduced. Growth took place from February to November. During this period temperature ranged from ?1 to 16°C, the average photon flux density (ifI) (per 24 h) from 9 to 570 μE · m^?2 · s^?1 (0.5 m depth), and the length of the days (I > 1 μE · m^?2 · s^?1) from 6 to 24 h. Light-limited growth was evident when the product of the average daily light and the chlorophyll/N ratio was < 10; this occurred mostly in early spring and late autumn. Peak densities (> 800 for the Thalassiosira spp. and > 1300–1400 μE · m^?2 · s^?1 for Skeletonema) seem to inhibit growth. The highest rates recorded were ≈1.6 doubl. · day^?1 (July, 15–16°C).The three species exhibit different ecological behaviour. Skeletonema is eurythermal (Q₁₀ = 1.8), whereas Thalassiosira pseudonana favours high temperatures, and T. gravida temperatures < 10°C. Moreover, Skeletonema has generally less chlorophyll and more phosphorus and ATP (≈ 1.4 ×) than the other two species. In Skeletonema, the ATP level seems related to the light-governed growth rate, and independent of temperature. In Thalassiosira no such correlation was found.     

Parameters of photosynthesis: definitions, theory and interpretation of results

Regrettably, parentheses are missing in Equation 16 on p. 1649.Equation 16 should read:      

BIO-OPTICAL CHARACTERISTICS AND PHOTOADAPTIVE RESPONSES IN THE TOXIC AND BLOOM-FORMING DINOFLAGELLATES GYRODINIUM AUREOLUM,GYMNODINIUM GALATHEANUM,AND TWO STRAINS OF PROROCENTRUM MINIMUM1

Photoadaptive responses in the toxic and bloom-forming dinoflagellates Gyrodinium aureolum Hulbert, Gymnodinium galatheanum Braarud, and two strains of Prorocentrum minimum (Pavillard)Schiller were evaluated with respect to pigment composition, light-harvesting characteristics, carbon and nitrogen contents, and growth rates in shade- and light-adapted cells. The two former species were grown at scalar irradiances of 30 and 170 μmol · m ^?2 at a 12-h daylength at 20° C. The two strains of P. minimum were grown at 35 and 500 μmol. m^?2· s^?1 at a 2-h daylength at 20° C. For the first time, chlorophyll (chl) c₃, characteristic of several bloom-forming prymnesiophytes, was detected in G. aureolum and G. galatheanum. Photoadaptional status affected the pigment composition strongly, and the interpretation of the variation depended on whether the pigment composition was normalized per cell, carbon, or chl a. Species-specific and photoadaptional differences in chl a-specific absorption (°a_c, 400–700 nm) and chl a-normalized fluorescence excitation spectra of photosystem II fluorescence with or without addition of DCMU (°F and °F_DCMU 400–700 nm) were evident. Gyrodinium aureolum and G. galatheanum exhibited in vivo spectral characteristics similar to chl c₃-containing prymnesiophytes in accordance with their similar pigmentation. Prorocentrum minimum had in vivo absorption and fluorescence characteristics typical for peridinin-containing dinoflagellates. Species-specific differences in in vivo absorption were also observed as a function of package effect vs. growth irradiance. This effect could be explained by differences in intracellular pigment content, cell size/shape, and chloroplast morphology/numbers. Light- and shade-adapted cells of P. minimum contained 43 and 17% of photoprotective carotenoids (diadino + diatoxanthin) relative to chl a, respectively. The photoprotective function of these carotenoids was clearly observed as a reduction in °F and °F ^DCMU at 400–540 nm compared to °ac in light-adapted cells of P. minimum. Spectrally weighted light absorption (normalized to chl a and carbon, 400–700 nm) varied with species and growth conditions. The use of quantum-corrected and normalized fluorescence excitation spectra with or without DCMU-treated cells to estimate photosynthetically usable light is discussed. The usefulness of in vitro absorption and fluorescence excitation spectra for estimation of the degradation status of chl a and the ratio of chl a to total pigments is also discussed.     

Aspects of statin prescribing in Norwegian counties with high,average and low statin consumption – an individual-level prescription database study

Background  

A previous study has shown that variations in threshold and intensity (lipid goal attainment) of statins for primary prevention contribute to regional differences in overall consumption of statins in Norway. Our objective was to explore how differences in prevalences of use, dosing characteristics, choice of statin and continuity of therapy in individual patients adds new information to previous results.     

Effect of light regime upon growth rate and chemical composition of a clone of Skeletonema costatum from the Trondheimsfjord, Norway

Nutrient-saturated cultures of Skeletonema costatum were grownat 15C and 42 combinations of photon flux density (PFD) anddaylength. The growth rate increased with the daylength andPFD up to 460–630 µE m^–2 s^–1 (maximum2.5 doublings day At 2000 µE m^–2 s^–1 the growthrate was reduced by 45%. The chlorophyll (chl) content of thecells and the rate of production of carbon per unit chlorophylland ambient light increased for declining light regimes as didcellular nitrogen and carbon. The N/C ratio, cellular phosphorusand ratios between in vivo fluorescence, with and without DCMU,and chlorophyll varied negligibly. The ATP/C ratio was linearlyrelated to the growth rate. The results were described mathematically.The chl/C ratio was low both in strong light and in marginallylow light, corresponding to low cellular chlorophyll and highcellular carbon, respectively. The observed increase in cellularnitrogen and carbon at shade adaptation probably represent anincrease in the size of internal stores of organic nitrogenand may imply that Skeletonema cells become enriched with organicnitrogen when staying in nitrate-rich subsurface layers, e.g.in or below a nutricline. However, close to zero growth in marginallight the cells become greatly enriched with respect to everymeasured factor. Such cells may be physiologically resting stageswhich may ensure survival during dark periods and promote rapiddevelopment during the initial phase of blooms. Cultures andnatural blooms of Skeleronema in the Trondheimsfjord exhibitvery similar patterns of variation.     

Nutrient status of phytoplankton communities in Norwegian waters (marine, brackish, and fresh) as revealed by their chemical composition

The chemical composition of nutrient-saturated cultures of Emilianiahuxleyi, Amphidinium carterae, and Staurastrum luetkemuelleriwas studied. The variation in chemical composition of naturalphytoplankton communities in the North Sea, the Trondheimsfjord,and a eutrophic lake was also studied. Nutrient status was evaluatedby measurement of the algal protein/carbohydrate, N/C, P/C,and N/P ratios. Tests for P-deficiency were carried out by measuringthe increase in ATP upon addition of phosphate. At saturationthe N/C ratio was {small tilde}0.14 in marine species and {smalltilde}0.05 in Staurastrum. Saturation P/C ratios (excludingpolyphosphates) were species-dependent, ranging from 0.017 (Skeletonema)to 0.006 (Amphidinium). Amphidinium and Staurastrum store polyphosphateswhen grown in P-rich media; true marine planktonic species donot. Natural communities tended to be close to nutrient saturationat low biomass densities and nutrient deficient at high densities.In the North Sea, nitrogen was clearly limiting. In waters offthe Møre coast and in the Trondheimsfjord, growth wasnearly balanced with respect to N and P at high salinities (>25)and clearly P-limited in brackish fjord waters. In dense communities,the N/P ratio was inversely related to salinity. Freshwatercommunities were clearly P-limited, but responses were dampenedwhen daphnia or whitefish were introduced, due to increasedexcretion of nutrients. ¹Contribution No. 212, Trondheim Biological Station, N-7001Trondheim, Norway.     

Studies on the phytoplankton ecology of the Trondheimsfjord. I. The chemical composition of phytoplankton populations

Samples of phytoplankton populations from the Trondheimsfjord, collected in 1970 and the first five months of 1971, have been analysed for carbohydrate, protein, lipid, and phosphorus. Lipid was in all cases less than 10% of the organic dry matter. The $\text{N}\text{P}$ ratio was remarkably constant, but the ratio protein/carbohydrate varied between wide limits. For samples consisting mainly of dinoflagellates, the protein/carbohydrate ratio was always low, due to a large amount of insoluble polysaccharides, probably corresponding to material in the cell walls.For diatoms, the carbohydrates may conveniently be divided into three fractions: 1) an acidsoluble glucan of the β-1, 3-linked type; 2) an alkali-soluble fraction giving a complex mixture of monosaccharides on hydrolysis and, 3) an insoluble glucan. The amounts of acid-soluble glucan varied from 7.7 to 36.5% of organic dry matter and these changes are the main cause of the variation of the protein/carbohydrate ratio of diatom samples. For diatom samples this ratio is a valuable indicator of the physiological state of the population. The variations observed in this study are discussed.     

Biooptical characteristics of PSII and PSI in 33 species (13 pigment groups) of marine phytoplankton,and the relevance for pulse‐amplitude‐modulated and fast‐repetition‐rate fluorometry1

We studied the variability of in vivo absorption coefficients and PSII‐scaled fluorescence excitation (fl‐ex) spectra of high light (HL) and low light (LL) acclimated cultures of 33 phytoplankton species that belonged to 13 different pigment groups (PGs) and 10 different phytoplankton classes. By scaling fl‐ex spectra to the corresponding absorption spectra by matching them in the 540–650 nm range, we obtained estimates for the fraction of total chl a that resided in PSII, the absorption of light by PSII, PSI, and photoprotective carotenoids. The in vivo red peak absorption maxima ranged from 673 to 679 nm, reflecting bonding of chl a to different pigment proteins. A simple approach is presented for quantifying intracellular self‐shading and evaluating the impact of photoacclimation on biooptical characteristics of the different PGs examined. In view of these results, parameters used in the calculation of oxygenic photosynthesis based on pulse‐amplitude‐modulated (PAM) and fast‐repetition‐rate (FRR) fluorometers are discussed, showing that the ratio between light available to PSII and total absorption, essential for the calculation of the oxygen release rate (using the PSII‐scaled fluorescence spectrum as a proxy) was dependent on species and photoacclimation state. Three subgroups of chromophytes exhibited 70%–80%, 60%–80%, and 50%–60% chl a in PSII‐LHCII; the two subgroups of chlorophytes, 70% or 80%; and cyanobacteria, only 12%. In contrast, the mean fraction for chromo‐ and chlorophytes of quanta absorbed by PSII was 73% in LL‐ and 55% in HL‐acclimated cells; thus, the corresponding ratios 0.55 and 0.73 might be used as correction factors adjusting for quanta absorbed by PSII for PAM and FRR measurements.     

THE SIGNIFICANCE OF INTRACELLULAR SELF-SHADING ON THE BIOOPTICAL PROPERTIES OF BROWN,RED, AND GREEN MACROALGAE1

The spectral light absorption and fluorescence excitation spectra from thalli (in vivo) and thylakoid micelles (no intracellular self-shading) from the macroalgae Laminaria digitata (L.) Edmonson, Palmaria palmata (L.) Kuntze, and Ulva sp. were examined. In all the examined species, the intracellular self-shading (i.e. the package effect) was highly significant to the extent that absorption peaks related to individual pigments were significantly suppressed. Thus, for studies related to taxonomy and physiology, “unpacked”spectra may be more informative. Thylakoid micelle preparations offer opportunities to examine more closely species-specific differences in the packaging effect, light harvesting and utilization, and pigmentation. As well, the thylakoid micelles, when analyzed with biochemical assays or fluoromtry, may provide insight into components of the photosynthetic machinery that are specific to different classes of mncroalgne. We present the Chl a-specific absorption coefficients and the corresponding fluorescence excitation spectra of macroalgal thylakoid micelles for the three species.     

RATE OF O2 PRODUCTION DERIVED FROM PULSE‐AMPLITUDE‐MODULATED FLUORESCENCE: TESTING THREE BIOOPTICAL APPROACHES AGAINST MEASURED O2‐PRODUCTION RATE1

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.