Short-term variability of photosynthesis in natural marine phytoplankton populations

Harmonic regression analysis has been used to determine the short-term variability in the photosynthetic rate (mgC/mg chlorophyll a/h) of phytoplankton in three inlets of Japan. In natural water without large zooplankton present, the photosynthetic rate [log P=log (100xmgC/mg chlorophyll a/h)] can be expressed as (B+A cos T). Factor B represents the average photosynthetic rate, of which the maximum is usually designated as P _max, and Factor A corresponds to the slope of the regression line. The phase of the periodicity, represented by T, is adjusted to give the highest correlation: usually T is expressed as [360/24 x (local time + 4)] in degrees. The correlation between Factors A and B is very high (r=0.95, P<0.001), indicating that Factor A may depend upon Factor B (potential activity of chlorophyll a). Both Factors A and B decrease with decreasing irradiance, but the slope of each regression between Factor A and irradiance varies with season. Continuous darkness reduces the phase of the periodicity to one cycle a day when phytoplankton has multiple cycles of photosynthetic rate per day. Adequate nutrient supply from zooplankton regeneration may cause an increase in Factor B; however, excess density of zooplankton decreases Factor A.     

The phenology of phytoplankton blooms: Ecosystem indicators from remote sensing

Remote sensing offers many advantages in the development of ecosystem indicators for the pelagic zone of the ocean. Particularly suitable in this context are the indicators arising from time series that can be constructed from remotely sensed data. For example, using ocean-colour radiometry, the phenology of phytoplankton blooms can be assessed. Metrics defined in this way show promise as informative indicators for the entire pelagic ecosystem. A simple phytoplankton–substrate model, with forcing dependent on latitude and day number is used to explore the qualitative features of bloom phenology for comparison with the results observed in a suite of 10-year time series of chlorophyll concentration, as assessed by remote sensing, from the Northwest Atlantic Ocean. The model reveals features of the dynamics that might otherwise have been overlooked in evaluation of the observational data.     

Simultaneous nitrogen and silicate deficiency of a phytoplankton community in a coastal jet-front

The nutritional status of a phytoplankton community was investigated in a coastal jet-front located in the Gulf of St. Lawrence, Canada, in 1987. During the sampling period, the frontal community was mainly composed of the diatomsChaetoceros debilis, Skeletonema costatum, Thalassiosira gravida andC. pelagicus. As previously reported for the St. Lawrence, some frontal stations were depleted both in nitrate and silicate. At stations impoverished in nitrate, internal nitrate pool concentrations were low or undetectable, suggesting that cells had not, recently, been exposed to a nitrate flux which exceeded the nitrate assimilation rate. At these impoverished stations, however, ambient and intracellular concentrations of ammonium and urea were high, suggesting that the community was not nitrogen-deficient. The comparison between the ambient silicate concentrations and the silicate requirement (K _s ) of the dominant diatoms suggests thatC. debilis andS. costatum were Si-deficient. This is further supported by the low silicate uptake rates and intracellular concentrations measured at the silicate impoverished stations. The silicate deficiency also resulted in a decrease in the seston and phytoplankton N:C ratios.Please address all correspondence and requests for reprints to Dr Levasseur at his new address: Maurice Lamontagne Institute, 850 Route de la Mer, Mont-Joli, Québec G5H 3Z4, Canada     

Short-term variations in the pigment composition of a spring phytoplankton bloom from an enclosed experimental ecosystem

Phytoplankton samples taken during the spring bloom in the experimental enclosed ecosystem bags at Loch Ewe, Scotland, during 1983 were analysed for carotenoids and chlorophyll compounds using high-performance liquidchromatography (HPLC). Changes in the relative proportions of these pigments were related to day-to-day changes in the composition of the bloom and the physiological state of the algae. There is clear evidence for a change in the chlorophyllide a:chlorophyll a ratio, which reached a maximum as nutrient limitation occurred. No major qualitative changes in the carotenoid components were seen during the bloom; the relative proportion, however, of some carotenoids does provide useful information on the relative abundance of certain algal type in the phytoplankton.     

A spatial model of phytoplankton patchiness

The one-dimensional theory of critical-length scales of phytoplankton patchiness is developed to include phytoplankton growth and herbivore grazing as functions of time and space. The critical-length scale L _c for the pathch is then determined by the initial spatial distribution and concentration of the limiting nutrient and herbivores in addition to the daily averaged values of the growth and loss processes. The response of an initial phytoplankton patch to the stresses of turbulent diffusion, nutrient depletion, light periodicity, and nocturnal or continuous herbivore grazing is investigated numerically for several oceanic conditions. Nocturnal grazing, while less stressful on primary production than continous grazing, results in lower phytoplankton standing stocks. Increase in biomass of vertically migrating zooplankton results in a net loss of nutrient which might otherwise be egested, recycled, and utilized in the euphotic zone under continuous grazing conditions. The Ivlev constant is shown via sensitivity analysis to be a significant parameter ultimately influencing phytoplankton production. It is demonstrated numerically that diffusion of phytoplankton cells from areas of high concentration to low concentration prevents the local extinction of the standing stock, thereby rendering a positive herbivore grazing-threshold unnecessary for ecosystem stability.     

Developing an empirical model of phytoplankton primary production: a neural network case study

We describe the development of a neural network model for estimating primary production of phytoplankton. Data from an enriched estuary in the eastern United States, Chesapeake Bay, were used to train, validate and test the model. Two error backpropagation multilayer perceptrons were trained: a simpler one (3-5-1) and a more complex one (12-5-1). Both neural networks outperformed conventional empirical models, even though only the latter, which exploits a larger suite of predictive variables, provided truly accurate outputs. The application of this neural network model is thoroughly discussed and the results of a sensitivity analysis are also presented.     

Two roads to two sexes: unifying gamete competition and gamete limitation in a single model of anisogamy evolution

Recent studies have revealed the importance of self-consistency in evolutionary models, particularly in the context of male–female interactions. This has been largely ignored in models of the ancestral divergence of the sexes, i.e., the evolution of anisogamy. Here, we model the evolution of anisogamy in a Fisher-consistent context, explicitly taking into account the number of interacting individuals in a typical reproductive group. We reveal an interaction between the number of adult individuals in the local mating group and the selection pressures responsible for the divergence of the sexes. The same underlying model can produce anisogamy in two different ways. Gamete competition can lead to anisogamy when it is relatively easy for gametes to find each other, but when this is more difficult and gamete competition is absent, gamete limitation can provide another route for anisogamy to evolve. In line with earlier models, organismal complexity favors anisogamy. We argue that the early contributions of Kalmus and Scudo, largely dismissed as group selectionist, are valid under certain conditions. Linking their work with the contributions of Parker helps to explain why precisely males keep producing more sperm than can ever lead to offspring: sperm could evolve to provision zygotes but this brings little profit for the effort required, because sperm would have to be equipped with provisioning ability before it is known which sperm will make it to the fertilization stage. This insight creates a logical link between paternal care under uncertain paternity (where again investment is selected against when some investment never brings about genetic benefits) and gamete size evolution.     

Carbon fixation in marine phytoplankton: carboxylase activities and stable carbon-isotope ratios; physiological and paleoclimatological aspects

We measured the activity of three carboxylases: RuBP carboxylase, PEP carboxylase and PEP carboxykinase of marine phytoplankton species in culture and in natural communities. Activities of the three carboxylases were measured simultaneously with stable carbon-isotope ratios. The enzymatic activities have been used to estimate the importance of carboxylation and its impact on the ¹³C:¹²C ratio (expressed as ¹³C). The marine phytoplankton species in culture were Fragilariopsis kerguelensis, Nitzschia turgiduloides, Skeletonema costatum, Phaeodactylum tricornutum, Isochrysis galbana, Dunaliella marina, and Prorocentrum micans, and the field samples were collected from different depths off the coast of Portugal (August/September, 1981). Our results indicate that, as in terrestrial plants, the ¹³C value is a good indicator of the extent of carboxylation. RuBP carboxylase activity was always predominant, whereas the ¹³C value never reached values typical of the C₄ pathway. The carboxylases could be PEP carboxylase (in dinoflagellates) or PEP carboxykinase (in diatoms). carboxylation increased at the end of the exponential growth phase in a diatom culture and with increased biomass in natural samples. We interpret these increases as an adaptative response mechanism to poor environmental conditions, especially to low light intensity.     

Response of marine phytoplankton to nitrogen deficiency: Decreased nitrate uptake vs enhanced ammonium uptake

The uptake of nitrate and ammonium was measured separately in uni-algal, nitrogen-deficient cultures of four species of marine phytoplankton. Nitrogen-deficient phytoplankton took up ammonium at initial rates which greatly exceeded those measured for nitrogen-sufficient phytoplankton. However, nitrate uptake by nitrogendeficient cultures was generally much slower than either nitrate or ammonium uptake by nitrogen-sufficient cultures or ammonium uptake by nitrogen-deficient cultures. Considerable species differences were observed in the degree to which nitrogen deficiency increased ammonium uptake or decreased nitrate uptake. Loss of ability to take up nitrate, but enhanced ability to take up ammonium, as a result of nitrogen deficiency may be an adaptation to the different mechanisms by which nitrate and ammonium are supplied to the euphotic zone. In areas with an intermittent supply of nitrogen, changes in the ability of some species to take up nitrogen as a result of nitrogen starvation will influence species composition and complicate interpretations of measurements of nitrogen uptake.Contribution no. 1249 from the Department of Oceanography, University of Washington, and contribution no. 82006 from the Bigelow Laboratory for Ocean Sciences     

Isotopic composition (C & N) of the suspended particles and N uptake by phytoplankton in a shallow tropical coastal lagoon

The spatial distribution of the C/N ratios and variations in δ¹³C and δ¹⁵N of suspended particulate matter were used to characterise their source in Asia’s largest brackish water lagoon, Chilika, India. In addition, the significance of re-mineralised nutrients in the primary productivity of the shallow lagoon was also determined through quantification of the subsurface nitrogen uptake conditions at two relatively stable locations in the lagoon. The results indicated that the influence of terrestrial organic matter was the maximum in the northern sector and was relatively limited at the central and southern part of the lagoon. In situ ¹⁵N uptake experiments (daytime) under biogeochemically stable conditions revealed that the N uptake by phytoplankton ranged between 0.24 and 1.01?mM?m^?3?h^?1 during pre-monsoon and post-monsoon seasons. New production and regenerated production in the shallow lagoon was also estimated by calculating f-ratios (ratio of nitrate assimilation by phytoplankton to total nitrogenous nutrient assimilation, have been estimated), which varied from 0.52 in the post-monsoon to 0.38 in the pre-monsoon. Lowering of the f-ratio from post- to pre-monsoon indicated a dominance of mineralisation over the new production.     

Short-term variability and control of phytoplankton photosynthetic activity in a macrotidal ecosystem (the Strait of Dover, eastern English Channel)

Short-term changes in phytoplankton photosynthetic activity were studied during different periods of the years 2009 and 2010 in the coastal waters of a macrotidal ecosystem (the Strait of Dover, eastern English Channel). During each sampling period, samples were taken every 1.45 h., from sunrise to sunset, during at least 5 days distributed along a complete spring–neap tide cycle. The photosynthetic parameters were obtained by measuring rapid light curves using pulse amplitude modulated fluorometry and were related to environmental conditions and phytoplankton taxonomic composition. The maximum quantum yield (F _v/F _m) showed clear light-dependent changes and could vary from physiological maxima (0.68–0.60) to values close to 0.30 during the course of 1 day, suggesting the operation of photoprotective mechanisms. The maximum electron transport rate (ETR_m) and maximal light utilization efficiency (α) were generally positively correlated and showed large diel variability. These parameters fluctuated significantly from hour to hour within each day and the intraday pattern of variation changed significantly among days of each sampling period. Stepwise multiple linear regressions analyses indicated that light fluctuations explained a part of this variability but a great part of variability stayed unexplained. F _v/F _m, ETR_m and α were not only dependent on the light conditions of the sampling day but also on those of the previous days. A time lag of 3 days in the effect of light on ETR_m and α variation was highlighted. At these time scales, changes in phytoplankton community structure seemed to have a low importance in the variability in photosynthetic parameters. The photoacclimation index E _k showed a lower variability and was generally different from the incident irradiance, indicating a limited acclimation capacity with a poor optimization of light harvesting during the day. However, in well-mixed systems such as the Strait of Dover, the short-term photoacclimation is disrupted by the high level of variability in environmental conditions. Also, the variability observed in the present study can be associated with a particular kind of photosynthetic response: the “E _k-independent” variability. The physiological basis of this photosynthetic response is largely unresolved and further researches on this subject are still required to better explain the dynamics of phytoplankton activity in the Strait of Dover.     

West Indian Ocean phytoplankton: a numerical investigation of phytohydrographic regions and their characteristic phytoplankton associations

Phytoplankton samples were collected from a large area of the West Indian Ocean and 237 different species identified. The object of this study was to define structure in the species distributions and to relate this to environmental factors. Numerical methods of cluster analysis are described, with adaptation and application to this ecological study of phytoplankton. The phytoplankton stations were grouped according to their phytoplankton populations; these groupings were related firstly to the monsoon seasons, and secondly to the hydrology, in order to derive phytohydrographic regions. The species were then grouped according to hydrographical distribution in order to derive the floral elements (species associations) which typify the different regions. The samples divided seasonally into 2 overlapping clusters, those sampled prior to the South West Monsoon, and those sampled subsequently. In both seasonal groups 4 main phytohydrographic regions were isolated from (1) equatorial subsurface water; (2) the equatorial undercurrent boundary; (3) the South Equatorial Current; (4) the surface tropical water. A total of 11 different floral elements were derived to account for the phytohydrographic groupings. The distribution of the largest element (50 species) was centred in equatorial subsurface water. This element dominated all samples, and was considered to comprise the endemic Indian Ocean flora. The effects of the other minor floral elements were superimposed on the dominance or otherwise of this element in the samples. These different minor floral elements were characteristic of (a) different seasonal divisions; (b) regional differences in equatorial subsurface water; (c) traversing currents; (d) nutrientrich regions of instability.     

Nutrient-limited toxin production and the dynamics of two phytoplankton in culture media: A mathematical model

In this paper we have proposed and analyzed a simple mathematical model consisting of four variables, viz., nutrient concentration, toxin producing phytoplankton (TPP), non-toxic phytoplankton (NTP), and toxin concentration. Limitation in the concentration of the extracellular nutrient has been incorporated as an environmental stress condition for the plankton population, and the liberation of toxic chemicals has been described by a monotonic function of extracellular nutrient. The model is analyzed and simulated to reproduce the experimental findings of Graneli and Johansson [Graneli, E., Johansson, N., 2003. Increase in the production of allelopathic Prymnesium parvum cells grown under N- or P-deficient conditions. Harmful Algae 2, 135–145]. The robustness of the numerical experiments are tested by a formal parameter sensitivity analysis. As the first theoretical model consistent with the experiment of Graneli and Johansson (2003), our results demonstrate that, when nutrient-deficient conditions are favorable for the TPP population to release toxic chemicals, the TPP species control the bloom of other phytoplankton species which are non-toxic. Consistent with the observations made by Graneli and Johansson (2003), our model overcomes the limitation of not incorporating the effect of nutrient-limited toxic production in several other models developed on plankton dynamics.     

Turbulence and phytoplankton diversity: A general model of the “paradox of plankton”

The principle of “competitive exclusion” predicts that no two species can occupy the same ecological niche at the same time and place (Hardin, 1960). Hutchinson (1953, 1961) suggested that the vast diversity of phytoplankton observed in many aquatic environments presents an apparent contradiction to this principle. Since all phytoplankton compete for the same basic resources, and since the euphotic zones of most natural waters are relatively homogeneous, such coexisting plankters appear to be simultaneously occupying the same niche. In this paper we present simulation results from a mathematical model wherein we examine the hypothesis that physical turbulence in an aquatic system can mollify interactive pressures between plankton populations and allow coexistence of species competing for the same resources. Using Bella's (1972) highly simplified model as a point of departure, we develop a new model, explicitly incorporating gross physiological mechanisms, to investigate the effects of both advective and turbulent components of water movement on the growth of three competing phytoplankton species. We observed that, in the absence of water motion, no two species were able to coexist, while under the hypothetical conditions of advection without turbulence (laminar flow), just two species were able to occur contemporaneously. Coexistence of all three species was achieved only with the addition of a random turbulent component to the model's hydrodynamic function. Moreover, this general coexistence was observed only when the major turbulent frequency approached the turnover rate of phytoplankton populations. We suggest that there is a limited region of periodicities and magnitudes for hydrodynamic energy in which phytoplankton can coexist, and that most natural aquatic environments fall within this region. We further speculate that, in general, the coupling of physical and biological processes in nature may be influenced by the relative frequency characteristics of those processes.     

Simulation model for wind-driven summer phytoplankton dynamics in Auke Bay,Alaska

A model was developed for use in simulating effects of short-term wind-mixing of the water column on estuarine phytoplankton dynamics. Simulation results, using parameter values estimated from field data, exhibited maxima observed in phytoplankton standing crop over a summer in Auke Bay, Alaska. Short-term wind-mixing of the water column can be important in determining the time of occurrence of phytoplankton community productivity pulses in deep estuaries.     

A three-component model of phytoplankton size class for the Atlantic Ocean

A three-component model was developed which calculates the fractional contributions of three phytoplankton size classes (micro-, nano- and picoplankton) to the overall chlorophyll-a concentration in the Atlantic Ocean. The model is an extension of the Sathyendranath et al. (2001) approach, based on the assumption that small cells dominate at low chlorophyll-a concentrations and large cells at high chlorophyll-a concentrations. Diagnostic pigments were used to infer cell size using an established technique adapted to account for small picoeukaroytes in ultra-oligotrophic environments. Atlantic Meridional Transect (AMT) pigment data taken between 1997 and 2004 were split into two datasets; 1935 measurements were used to parameterise the model, and a further 241 surface measurements, spatially and temporally matched to chlorophyll-a derived from SeaWiFS satellite data, were set aside to validate the model. Comparison with an independent global pigment dataset (256 measurements) also supports the broader-scale application of the model. The effect of optical depth on the model parameters was also investigated and explicitly incorporated into the model. It is envisaged that future applications would include validating multi-plankton biogeochemical models and improving primary-production estimates by accounting for community composition.     

硫脲用量对菠菜生理指标及土壤氮素转化的影响

采用盆栽法,研究了硫脲不同用量(0%、1%、3%、5%、7%)对菠菜生长、硝酸盐累积、土壤氮含量及叶绿体色素含量的影响.结果表明,不同硫脲用量对菠菜的单盆重、株高均有提高.随硫脲用量的增加,菠菜叶绿体色素含量均有提高;硫脲能显著降低菠菜硝酸盐含量,随着硫脲用量的增加,降低效果提高,其中硫脲用量为7%时,降低效果达极显著水平.硫脲对土壤硝态氮累积的抑制作用随硫脲用量的增加而增强,硫脲用量达7%时,抑制作用表现为极显著.     

Nitrate storage by phytoplankton in a coastal upwelling environment

The storage of nitrate by phytoplankton cells during the early phases of upwelling was studied in coastal stations off northern Spain (southern Bay of Biscay) between 1990 and 1994. In this region, a persistent upwelling during summer is characterised by intermittent pulses of variable intensity, and increased nutrient concentrations in the surface layer. The main effect of an upwelling pulse on phytoplankton distribution is the shifting of the chlorophyll a and primary production maxima to near the surface. When the upwelling relaxes, thermal stratification of the water column occurs, and a distinct subsurface chlorophyll maximum develops below the production maximum. An accumulation of intracellular nitrate characterized the early phases of upwelling (mean = 2.73 μmol N m⁻³), maximum concentrations being attained at depths where biomass and production values were moderate. In contrast, phytoplankton cells from non-upwelling situations contained significantly lower concentrations of intracellular nitrate (mean = 0.17 μmol N m⁻³). The variations in the intracellular pool of nitrate may result from the differential allocation of resources within the cell as a result of variations in the energy available, since the uptake and assimilation of nitrate is a relatively expensive process involving several enzymatic systems. We hypothesize that nitrate storage by phytoplankton cells is characteristic of early phases of upwelling and is linked to patterns of carbon fixation. Average nitrogen budgets for upwelling and non-upwelling situations indicate that intracellular nitrate reserves are not responsible for maintaining high phytoplankton growth rates, since they only account for <2% of daily primary production during upwelling events. Received: 28 August 1996 / Accepted 3 December 1996