Seasonal variations of phytoplankton photosynthate partitioning in two lakes of different trophic level

The aim of this study was to compare vertical and seasonal variationsof inorganic carbon allocation into macromolecules by the phytoplanktonpopulation in a eutrophic lake (Lake Aydat) and an oligo-mesotrophiclake (Lake Pavin). Biochemical fractionation was conducted byconsecu tive differential extractions in order to separate proteins,polysaccharides, lipids, and low molecular weight compounds(LMW). The ratio of light absorption at480 and 665 nm by acetoneextracts of phytoplankton pigments was used as an indicatorof the nutritional statusof natural phytoplankton populations.Our results show that in Lake Aydat, the main photosyntheticend productswere poly saccharides, whereas in Lake Pavin, radioactivitywas predominantly incorporated into the protein fraction. Moreover,the seasonal cycles of mixing and stratification in these twolakes affected the pattern of ¹⁴C incorporation into LMW andmacromolecules. An increase in the relative synthesis of proteinsoccurred during stratification periods. It was linked to anincrease in temperature and nutrient limitation further complicatedby the shift in species composition of the populations. Differences recorded both between the two lakes of different trophicstatus and between seasons confirm that the proportion of carbonincorporated into proteins might be a useful indicator of thephysio logical status of phvtoplankton communities.     

Nutritional diagnosis of phytoplankton in Lake Baikal

To diagnose the nutritional status of phytoplankton in Lake Baikal, surveys for the determination of concentrations of particulate carbon (PC), nitrogen (PN) and phosphorus (PP) and their ratios were conducted at six stations in March, June, August and October 1999. The concentrations of PC and PN were lower than, and those of PP were similar to, those in another mesotrophic lake except at the station near the mouth of the largest input river, Selenga River, of Lake Baikal. The PC : PN : PP ratio was 102 : 13 : 1, considerably close to the Redfield ratio. The ratio was constant against spatiotemporal changes. These indicate that phytoplankton in Lake Baikal were exposed to no deficiency in nitrogen nor phosphorus. From a viewpoint of the nutritional status of phytoplankton, Lake Baikal might be viewed as an ocean rather than as a lake.     

Changes in alkaline phosphatase activity and phosphate uptake in P-limited phytoplankton,induced by light intensity and spectral quality

Alkaline phosphatase activity and P uptake were determined in P-limited Dunaliella tertiolecta, Thalassiosira pseudonana, Phaeodactylum tricornumtum, and Prymnesium parvum grown under different light intensities and colors. Both intracellular and extracellular enzyme activities varied with the intensity and quality of light in a species-specific manner. The spectral composition of the light also affected P uptake kinetics. No correlation was found between enzyme activity and V_max both within a species and for pooled data for all four species, indicating that the change in uptake kinetics and enzyme activity was not related to P limitation, but induced by the light conditions. Changes in the optimum N:P ratio induced by light were also not related to P uptake kinetics or enzyme activity. These data suggest that light conditions may in themselves have profound effects on species competition for limiting nutrients. Furthermore, since both alkaline phosphatase activity and P uptake were influenced by the prevailing light conditions we suggest that these parameters be used cautiously when determining the P nutritional status of phytoplankton in nature.Address for reprint requests     

The impact of phytoplankton on spectral water transparency in the Southern Ocean: implications for primary productivity

Spectral water transparency in the Northern Weddell Sea was studied during Austral spring. The depth of the 1-% surface irradiance level (euphotic depth) varied between 35 and 109 m and was strongly influenced by phytoplankton biomass. Secchi depths were non-linearly related to euphotic depth. In phytoplankton-poor water, the most penetrating spectral region was restricted to a relatively narrow waveband in the blue (488 nm), but the range was broader, between 488 and 525 nm when phytoplankton were abundant. Water transparency in the red spectral range was always low and only to a small extent affected by phytoplankton. Two independent procedures were used to quantify the impact of phytoplankton on spectral water transparency: (1) Regression analysis of spectral in situ vertical light attenuation coefficients in the sea, against coincident chlorophyll concentrations. This method gave chlorophyll-specific light attenuation coefficients; the y-intercept could be interpreted as a measure of light attenuation by pure water plus non-algal material. (2) Spectra of in vivo light absorption derived by spectroscopy, using phytoplankton enriched to varying degrees onto filters. Thus chlorophyll-specific absorption cross-sections were determined. Estimates obtained by both procedures were in close agreement. By integrating over the spectrum of underwater irradiance, in situ chlorophyll-specific absorption cross sections of phytoplankton suspensions, related to all photosynthetically active radiation, were calculated. Light absorption by phytoplankton for photosynthesis is accomplished mainly in the blue spectral range. Also dissolved and particulate organic matter contributed to the attenuation of blue light. Because in water poor in phytoplankton, underwater irradiance was progressively restricted to blue light, chlorophyll-specific absorption cross-sections of phytoplankton, averaged over the spectrum of photosynthetically active irradiance, increased with water depth. In water with elevated phytoplankton biomass, overall light attenuation was generally enhanced. However, because the spectral composition of underwater light changed relatively little with depth, except immediately below the water surface, light absorption cross-sections of phytoplankton changed little below 10 m depth. Vertical differences in the proportions of underwater light absorbed by the phytoplankton community here were mainly dependent on biomass variations. Because of the comparatively small attenuation of blue light by non-algal matter, the efficiency of light harvesting by phytoplankton at any given concentration of chlorophyll in Antractic waters is greater than in other marine regions. At the highest phytoplankton biomass observed by us, as much as 70% of underwater light was available for phytoplankton photosynthesis. When phytoplankton were scarce, <10% of underwater light was harvested by phytoplankton.Contribution within the European Polarstern Study (EPOS), supported by the Deutsche Forschungsgemeinschaft, Grant Ti 115/16-1 to MMT, the European Science Foundation, and by the Alfred Wegener Institut für Polar-und Meeresforschung, Bremerhaven     

Can heterotrophic bacteria be important to marine light absorption?

The contribution of heterotrophic bacteria to particulate lightabsorption in the ocean has been traditionally considered insignificantas compared to that of phytoplankton and detritus. This viewhas been based on the general preswnption that heterotrophicmarine bacteria do not contain pigments with significant absorptionin the visible spectral range. However, there exist heterotrophicbacteria that synthesize carotenoid pigments, and carotenoid-richstrains of bacteria have often been isolated from natural seawatersamples taken in the open and coastal ocean. Because carotenoidsabsorb strongly in the blue spectral region, the heterotrophicbacteria may contribute more to marine light absorption thanhas been assumed. In order to make preliminary assessment ofsuch a contribution, we measured the absorption of a strainof carotenoid-containing heterotrophic bacteria (CHB) grownin the laboratory under differing conditions of light and nutrientavailability. These measurements showed that absorption cross-sectionsof CHB in the blue could be at least twice, and possibly oneorder of magnitude, higher than those of non-pigmented heterotrophicbacteria (NHB). In addition, the absorption features of CHBwere conserved under the differing light and nutrient conditions.We conclude that the role of heterotrophic bacteria in marinelight absorption needs to be re-evaluated. This will requirefurther laboratory studies to quantify the absorption cross-sectionsof marine bacteria with improved accuracy, as well as the developmentof a technique for the recognition and enumeration of CHB cellsin the ocean.     

Critical conditions for phytoplankton blooms

We motivate and analyse a reaction—advection—diffusion model for the dynamics of a phytoplankton species. The reproductive rate of the phytoplankton is determined by the local light intensity. The light intensity decreases with depth due to absorption by water and phytoplankton. Phytoplankton is transported by turbulent diffusion in a water column of given depth. Furthermore, it might be sinking or buoyant depending on its specific density. Dimensional analysis allows the reduction of the full problem to a problem with four dimensionless parameters that is fully explored. We prove that the critical parameter regime for which a stationary phytoplankton bloom ceases to exist, can be analysed by a reduced linearized equation with particular boundary conditions. This problem is mapped exactly to a Bessel function problem, which is evaluated both numerically and by asymptotic expansions. A final transformation from dimensionless parameters back to laboratory parameters results in a complete set of predictions for the conditions that allow phytoplankton bloom development. Our results show that the conditions for phytoplankton bloom development can be captured by a critical depth, a compensation depth, and zero, one or two critical values of the vertical turbulent diffusion coefficient. These experimentally testable predictions take the form of similarity laws: every plankton—water—light-system characterized by the same dimensionless parameters will show the same dynamics.     

DIFFERENCES BETWEEN IN VIVO ABSORPTION AND FLUORESCENCE EXCITATION SPECTRA IN NATURAL SAMPLES OF PHYTOPLANKTON

The fluorescence excitation spectrum of live phytoplankton cells represents the portion of light absorbed that has been effectively transferred to chlorophyll a of photosystem II, whereas light absorbed by photoprotective pigments will not lead to fluorescence. Therefore, the in vivo fluorescence excitation spectrum of phytoplankton has been used as a proxy for the action spectrum of phytoplankton in computations of primary production in the ocean. The distribution of chlorophyll a between photosystems, as well as variations in the pathway of energy inside the photosynthetic membrane, can also influence the fluorescence excitation spectrum. In this study, we investigated the contribution of photoprotective pigments to the differences found between in vivo absorption and fluorescence excitation spectra of phytoplankton measured during two cruises: one from Las Islas Canarias to Nova Scotia and another in the Labrador Sea. A comparison of normalized fluorescence excitation and absorption spectra showed high variability in the difference between absorption and fluorescence in the blue region of the spectrum for samples from the two cruises. This difference was not entirely correlated with the concentration of photoprotective carotenoids. In this paper, results are interpreted in terms of differences in pigment composition and known patterns of energy distribution in the photosystems of different algal groups.     

COMBINATION OF FLOW CYTOMETRY AND SINGLE CELL ABSORPTION SPECTROSCOPY TO STUDY THE PHYTOPLANKTON STRUCTURE AND TO CALCULATE THE CHL A SPECIFIC ABSORPTION COEFFICIENTS AT THE TAXON LEVEL1

The phytoplankton community structure of a hypertrophic lake was quantitatively determined with the aid of flow cytometry. The flow cytometry signals were calibrated to obtain cell‐specific information, such as the chl a content and the biovolume per cell. The reliability of this method was tested with laboratory cultures. The results of the phytoplankton structure in a hypertrophic lake with respect to chl distribution in the different algal groups obtained by flow cytometry were compared with the results from HPLC pigment fingerprinting. Both methods yield the percentage contribution of the different algal groups to total chl a. The chl a specific absorption coefficient of the phytoplankton (a^*_Phy) was determined via visible (VIS) spectroscopy of samples taken from a hypertrophic lake (Auensee) in 2003. The results indicated that a^*_Phy of the total cell suspension is dependent on the phytoplankton structure as well as on environmental factors. The linear relationship between a^*_Phy at 675 nm and the product of the chl a content per cell and the biovolume offered the possibility to normalize phytoplankton absorption spectra to acquire the taxon‐specific a^*_Phy. The estimated a^*_Phy (675 nm) values were used to normalize single cell absorption spectra at this wavelength to obtain the a^*_Phy between 400 and 750 nm for representatives of the major algal groups. Our measurements show that the absorption coefficient for the whole phytoplankton community varies within the season. Finally, we used the a^*_Phy and the chl a distribution to calculate the light absorption of each algal group in the hypertrophic lake.     

Carbon allocation under light and nitrogen resource gradients in two model marine phytoplankton1

Marine phytoplankton have conserved elemental stoichiometry, but there can be significant deviations from this Redfield ratio. Moreover, phytoplankton allocate reduced carbon (C) to different biochemical pools based on nutritional status and light availability, adding complexity to this relationship. This allocation influences physiology, ecology, and biogeochemistry. Here, we present results on the physiological and biochemical properties of two evolutionarily distinct model marine phytoplankton, a diatom (cf. Staurosira sp. Ehrenberg) and a chlorophyte (Chlorella sp. M. Beijerinck) grown under light and nitrogen resource gradients to characterize how carbon is allocated under different energy and substrate conditions. We found that nitrogen (N)‐replete growth rate increased monotonically with light until it reached a threshold intensity (~200 μmol photons · m^?2 · s^?1). For Chlorella sp., the nitrogen quota (pg · μm^?3) was greatest below this threshold, beyond which it was reduced by the effect of N‐stress, while for Staurosira sp. there was no trend. Both species maintained constant maximum quantum yield of photosynthesis (mol C · mol photons^?1) over the range of light and N‐gradients studied (although each species used different photophysiological strategies). In both species, C:chl a (g · g^?1) increased as a function of light and N‐stress, while C:N (mol · mol^?1) and relative neutral lipid:C (rel. lipid · g^?1) were most strongly influenced by N‐stress above the threshold light intensity. These results demonstrated that the interaction of substrate (N‐availability) and energy gradients influenced C‐allocation, and that general patterns of biochemical responses may be conserved among phytoplankton; they provided a framework for predicting phytoplankton biochemical composition in ecological, biogeochemical, or biotechnological applications.     

DIEL VARIATIONS IN OPTICAL PROPERTIES OF MICROMONAS PUSILLA (PRASINOPHYCEAE)1

Micromonas pusilla (Butcher) Manton et Parke, a marine prasinophyte, was used to investigate how cell growth and division affect optical properties of phytoplankton over the light:dark cycle. Measurements were made of cell size and concentration, attenuation and absorption coefficients, flow cytometric forward and side light scattering and chl fluorescence, and chl and carbon content. The refractive index was derived from observations and Mie scattering theory. Diel variations occurred, with cells increasing in size, light scattering, and carbon content during daytime photosynthesis and decreasing during nighttime division. Cells averaged 1.6 μm in diameter and exhibited phased division, with 1.3 divisions per day. Scattering changes resulted primarily from changes in cell size and not refractive index; absorption changes were consistent with a negligible package effect. Measurements over the diel cycle suggest that in M. pusilla carbon‐specific attenuation varies with cell size, and this relationship appears to extend to other phytoplankton species. Because M. pusilla is one of the smallest eukaryotic phytoplankton and belongs to a common marine genus, these results will be useful for interpreting in situ light scattering variation. The relationship between forward light scattering (FLS) and volume over the diel cycle for M. pusilla was similar to that determined for a variety of phytoplankton species over a large size range. We propose a method to estimate cellular carbon content directly from FLS, which will improve our estimates of the contribution of different phytoplankton groups to productivity and total carbon content in the oceans.     

Comparison of Photoacclimation in Twelve Freshwater Photoautotrophs (Chlorophyte,Bacillaryophyte, Cryptophyte and Cyanophyte) Isolated from a Natural Community

Different representative of algae and cyanobacteria were isolated from a freshwater habitat and cultivated in laboratory to compare their photoacclimation capacity when exposed to a wide range of light intensity and to understand if this factor may modify natural community dominance. All species successfully acclimated to all light intensities and the response of phytoplankton to increased light intensity was similar and included a decrease of most photosynthetic pigments accompanied by an increase in photoprotective pigment content relative to Chl a. Most species also decreased their light absorption efficiency on a biovolume basis. This decrease not only resulted in a lower fraction of energy absorbed by the cell, but also to a lower transfer of energy to PSII and PSI. Furthermore, energy funnelled to PSII or PSI was also rearranged in favour of PSII. High light acclimated organisms also corresponded to high non-photochemical quenching and photosynthetic electron transport reduction state and to a low Φ''_M. Thus photoacclimation processes work toward reducing the excitation pressure in high light environment through a reduction of light absorption efficiency, but also by lowering conversion efficiency. Interestingly, all species of our study followed that tendency despite being of different functional groups (colonial, flagellated, different sizes) and of different phylogeny demonstrating the great plasticity and adaptation ability of freshwater phytoplankton to their light environment. These adjustments may explain the decoupling between growth rate and photosynthesis observed above photosynthesis light saturation point for all species. Even if some species did reach higher growth rate in our conditions and thus, should dominate in natural environment with respect to light intensity, we cannot exclude that other environmental factors also influence the population dynamic and make the outcome harder to predict.     

Light absorption by phytoplankton: development of a matching parameter for algal photosynthesis under different spectral regimes

A spectral matching parameter (absorption efficiency, A_e) wasdeveloped to quantify the relationship between the light absorptionspectra of phytoplankton communities and the spectral irradianceof their ambient light field. A_e was defined as the ratio betweenthe amount of radiation absorbed by the phytoplankton in situand the amount absorbed in a spectrally flat light regime. Thisapproach was applied to our measurements of spectral absorptionfor the phytoplankton communities in six lakes in High ArcticCanada that spanned a range of bio-optical conditions. A_e valueswere calculated for the light spectrum down through the watercolumn and for 11 types of artificial light source. Spectralmatching varied among lakes and with depth. There was a significantlinear relationship between the relative change in A_e with depthand the diffuse attenuation coefficient K_d (r² = 0.52, P = 0.012for K_d for the 400–700 nm waveband; r² = 0.78, P = 0.0003for K_d at 440 nm). The tabulated values for the matching parameterA_e allow the comparison of photosynthesis versus irradiance(P versus E) curves among studies using different light sources.A_e estimates also facilitate the evaluation of chromatic adaptationin natural waters, and the calculation of spectrally adjusted,in situ primary production down through a water column fromP versus E relationships under a single spectral regime.     

Winter severity determines functional trait composition of phytoplankton in seasonally ice‐covered lakes

How climate change will affect the community dynamics and functionality of lake ecosystems during winter is still little understood. This is also true for phytoplankton in seasonally ice‐covered temperate lakes which are particularly vulnerable to the presence or absence of ice. We examined changes in pelagic phytoplankton winter community structure in a north temperate lake (Müggelsee, Germany), covering 18 winters between 1995 and 2013. We tested how phytoplankton taxa composition varied along a winter‐severity gradient and to what extent winter severity shaped the functional trait composition of overwintering phytoplankton communities using multivariate statistical analyses and a functional trait‐based approach. We hypothesized that overwintering phytoplankton communities are dominated by taxa with trait combinations corresponding to the prevailing winter water column conditions, using ice thickness measurements as a winter‐severity indicator. Winter severity had little effect on univariate diversity indicators (taxon richness and evenness), but a strong relationship was found between the phytoplankton community structure and winter severity when taxon trait identity was taken into account. Species responses to winter severity were mediated by the key functional traits: motility, nutritional mode, and the ability to form resting stages. Accordingly, one or the other of two functional groups dominated the phytoplankton biomass during mild winters (i.e., thin or no ice cover; phototrophic taxa) or severe winters (i.e., thick ice cover; exclusively motile taxa). Based on predicted milder winters for temperate regions and a reduction in ice‐cover durations, phytoplankton communities during winter can be expected to comprise taxa that have a relative advantage when the water column is well mixed (i.e., need not be motile) and light is less limiting (i.e., need not be mixotrophic). A potential implication of this result is that winter severity promotes different communities at the vernal equinox, which may have different nutritional quality for the next trophic level and ecosystem‐scale effects.     

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.     

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.     

Allelopathic effect of the aquatic macrophyte, Stratiotes aloides, on natural phytoplankton

1. A survey of different Dutch Stratiotes stands showed that the density of phytoplankton (except cyanobacteria) was always higher outside S. aloides than between the rosettes of S. aloides. Analyses of water samples revealed that nutrient limitation was unlikely to have caused the lower phytoplankton biomass in the vicinity of S. aloides. 2. An in situ incubation experiment in the Danube Delta, Romania, indicated allelopathic activity against phytoplankton in S. aloides stands. The growth rate of natural phytoplankton populations exposed to water from S. aloides stands was significantly lower than that of populations that had not been in contact with S. aloides exudates. 3. A laboratory microcosm experiment showed a significantly lower phytoplankton biomass in treatments with S. aloides exudates. Nutrient concentrations and the light intensity were high enough that the lower phytoplankton biomass could not be explained by nutrient or light limitation.     

Unusual allometry between in situ growth of freshwater phytoplankton under static and fluctuating light environments: possible implications for dominance

The effects of fluctuating light fields on the growth of phytoplanktonare not well understood and conclusions in the literature havebeen equivocal. Most studies have examined responses such asproductivity and chlorophyll a content (laboratory culture andfield tests) or growth rates (laboratory culture tests). Inthis study we examined the in situ growth rates of differenttypes of phytoplankton within two natural populations. Comparisonswere made between populations grown in a static environment(suspended in a fixed position in the water column) and an equivalentpopulation moving through the water column simulating the mixingof entrained phytoplankton. Growth under fluctuating light fieldsin this experiment only significantly (P < 0.05) increasedthe growth of the diatom Skeletonema and decreased the growthof Anabaena circinalis, Microcystis aeruginosa and Scenedesmussp. All other phytoplankton, including the genera Nitzschia,Fragilaria and Dactylococcopsis, did not have growth rates thatwere significantly different between static and fluctuatinglight treatments. A general pattern where diatoms grew best,followed by chlorophytes with the toxicogenic cyanophytes M.aeruginosa and A. circinalis growing least well, was distinguishedunder fluctuating irradiance. This seems consistent with thecommon occurrence of these groups of phytoplankton in the naturalenvironment. The cyanophytes Dactylococcopsis and Aphanothecedid not follow this pattern, with the former growing betterunder fluctuating light and the latter exhibiting an unusualgrowth pattern where growth was higher under lower light intensities.     

A comparison of primary production measurements using two laboratory systems with differences in light quality

Primary production measurements were carried out simultaneously, using two laboratory systems with different light conditions: (1) a'classical' incubator and (2) a Laboratory Scale Enclosure. The model used for calculating primary production (STEELE, 1965) does not correct for spectral changes caused by high phytoplankton biomass. In the incubator, light of almost all wavelengths decreased more or less according to the attenuation of total PhAR in water. In the LSE, high absorption was found of the blue light and some of the red light, which was due to the high sestonic concentration. The Steele function provided a good fit for both sets of data. The depth integrated gross production values derived from the simultaneous measurements were not significantly different.     

Phytoplankton as a stimulus for spawning in three marine invertebrates

A distinct annual reproductive cycle with spring spawning was observed in Strongylocentrotus droebachiensis Müller, Tonicella lineata Wood and Tonicella insignis Reeve. This study gives evidence that the cue for spawning in these species is the spring phytoplankton bloom. In 1973 spawning occurred abruptly in early April at the time of the spring phytoplankton outburst, but in 1974 spawning was less abrupt corresponding to the slow development of the phytoplankton bloom in that year. Animals were collected prior to spawning and maintained in the laboratory under various temperature and light regimes: at 5 ° and 14 °C in darkness, and at 5 ° and 14 °C in light conditions similar to those in the field. These animals did not spawn when spawning occurred in the field, but animals returned to the field from the laboratory did spawn. In the laboratory a large proportion of animals spawned when they were exposed to phytoplankton collected with a 50μm mesh net. The results suggest that some substance bound to or released by phytoplankton stimulates spawning. For species with planktotrophic larvae the synchronization of spawning with the phytoplankton bloom increases the probability of favourable food and temperature conditions for the development of the larvae and juveniles.     

Erosion,phosphorus and phytoplankton response in rivers of South-Eastern Norway

The development of P fractions and phytoplankton was studied in three rivers with varying concentrations of seston.Less than 1% of the yearly TP transport may take place during periods with high algal biomass.The observation of a high growth rate of phytoplankton in the rivers coinciding with high concentrations of RP, low content of seston and high TP:Chl a ratio, indicate that the growth was often not P-limiting. During short periods with high phytoplankton biomass the ratio TP:Chl a may be low, indicating that a high fraction of TP was available.The content of P in soil samples and in samples with high seston content was about 0.1% of dry weight, and the algal availability of P often varied between 25 and 75% of TP for both types of samples.Decreasing biomass or low growth rates were observed at secchi depths less than 0.5 m and seston concentrations less than about 25 mg dry weight 1^–1. High flow rate also depressed the development of the total phytoplankton biomass. The assimilation of available P is incomplete under such conditions, i.e. under conditions of light limitation and high dilution rate.The availability of P for phytoplankton in rivers with different length, light conditions and stream velocity is discussed.