A simple model illustrating the role of turbulence on phytoplankton blooms

 The problem of the vertical distribution of phytoplankton is considered in the presence of gravitational settling, turbulent mixing, population growth due to cell division and a constant rate of loss due to predation and natural death. Nutrients are assumed to be plentiful so that the production rate depends only on the light available for photosynthesis. The non-linear saturation of plankton growth is modeled by allowing the attenuation rate of light to be a linear function of the plankton density. The turbulent diffusivity is assumed constant which corresponds to a mixed layer depth very much greater than the depth of light penetration (euphotic depth). It is shown that an exact analytical solution of this non-linear problem is possible for an idealized model in which the functional dependence of production on light intensity is assumed to be a step function. Non-zero solutions are shown to exist only if the parameters characterizing the system are above a certain critical curve in a two dimensional parameter space. Numerical simulations using functional forms of the production curve that resemble the measured photosynthetic response of plankton, show, that the qualitative behavior of the system is similar to that of the idealized model presented. Comparisons are made with other analytical approaches to the problem. Received: 14 March 2002 / Revised version: 27 September 2002 / Published online: 28 February 2003 Key words or phrases: Self-shading – Plankton – Nonlinear – Turbulence     

Dynamics of the ‘echo’ effect in a phytoplankton system with nitrogen fixation

Consideration of nitrogen fixation adds a positive nonlinear feedback to plankton ecosystem models. We investigate the consequences of this feedback for secondary phytoplankton blooms and the response of phytoplankton dynamics to physical forcing. The dynamics of phytoplankton, Trichodesmium (the nitrogen fixer), and nutrients is modeled with a system of three differential equations. The model includes two types of nonlinear interactions: the competition of phytoplankton and Trichodesmium for light, and the positive feedback resulting from Trichodesmium recycling. A typical simulation of the model in time, with forcing by a varying mixed-layer depth, reveals a clear successional sequence including a secondary or ‘echo’ bloom of the phytoplankton. We explain this sequence of events through the stability analysis of three different steady states of the model. Our analysis shows the existence of a critical biological parameter, the ratio of normalized growth rates, determining the occurrence of ‘echo’ blooms and the specific sequence of events following a physical perturbation. The interplay of positive and negative feedbacks appears essential to the timing and the type of events following such a perturbation.     

Composition changes of phototrophic microbial communities along the salinity gradient in the solar saltern evaporation ponds of Eilat,Israel

There are several conflicting hypothesis that deal with the influence of flooding in the natural river–floodplain systems. According to the Flood Pulse Concept, the flood pulses are not considered to be a disturbance, while some recent studies have proven that floods can be a disturbance factor of phytoplankton development. In order to test whether flooding acts as a disturbance factor in the shallow Danubian floodplain lake (Lake Sakadaš), phytoplankton dynamics was investigated during two different hydrological years—extremely dry (2003) without flooding and usually flooded (2004). A total of 18 phytoplankton functional groups were established. The sequence of phytoplankton seasonality can be summarized P/D → E (W1, W2) → C/P (only in potamophase) → S2/H1/S_N/S1 → W1/W2 → P/D. The canonical correspondence analysis (CCA) demonstrated that the water level was a significant environmental variable in 2004. Due to the higher total biomass of Bacillariophyceae established under potamophase conditions, floodings in the early spring seem to be a stimulating factor for phytoplankton development. On the other hand, the flood pulses in May and June had dilution effects on nutrients, so that a significantly lower phytoplankton biomass was established indicating that flooding pulses can be regarded as a disturbance event. Such conditions supported diatom development (D, P, C species) and prolonged its dominance in the total phytoplankton biomass. A long-lasting Cyanoprokaryota bloom (various filamentous species—S1, S2, S_N and H1 representatives) with very high biomass characterized the limnophase (dry conditions) in summer and autumn of both years. In-lake variables (lake morphology, internal loadings of nutrients from sediments, light conditions) seem to be important for the appearance of Cyanoprokaryota bloom. The equilibrium phase was found during the Cyanoprokaryota bloom only in the extremely dry year. This study showed that depending on the time scale occurrence, flood pulses can be a stimulating or a disturbance factor for phytoplankton development in Lake Sakadaš. Handling editor: J. Padisak     

Predicting the spatial dominance of phytoplankton in a light limited and incompletely mixed eutrophic water column using the PROTECH model

1. A phytoplankton community model [Phytoplankton RespOnses To Environmental CHange (PROTECH)] was used to examine the effect of a wide range of varied light intensities and mixed depths upon simulated phytoplankton populations. Two different column lengths of the simulated water body were examined (the upper 5 m and the whole 14.5 m water column) for each scenario.
2. The hypotheses tested were that: (i) under low light intensity and/or deep mixing the simulated community will be dominated by a phytoplankter with a low critical light intensity; (ii) at high light intensity and shallow mixing the simulated community will be dominated by small, fast-growing phytoplankters; (iii) under all conditions, except deep mixing, the largest proportion of phytoplankton biomass will be found near the surface.
3. It was found under most conditions that, although there was a bloom in the upper column (dominated by algae such as Chlorella , Ceratium or Rhodomonas ), the largest phytoplankton biomass in the water column was located 9 m below the surface and consisted of solely Asterionella . This bloom was missed by the 5-m samples. Thus, using the whole column sample lengths, hypothesis (i) was not rejected but hypotheses (ii) and (iii) were refuted.
4. The inclusion of specific movement characteristics of phytoplankton in the model allowed the possibility of the dominance of multiple spaces within the water column and should be included in any model-based investigation of this topic. Further, the results from the model suggest that a reduced depth of mixing creates greater environmental heterogeneity, allowing more species to persist.     

Seasonal variation of mixing depth and its influence on phytoplankton dynamics in the Zeya reservoir, China

In reservoirs or lakes, mixing depth affects growth and loss rates of phytoplankton populations. Based on 1-year data from the Zeya reservoir, China, we scaled the mixing depth throughout a whole year by utilizing cluster analysis, and then investigated its influence on phytoplankton dynamics and other physical and chemical parameters. Over the whole year, all physical and chemical parameters except TN and temperature had significant correlations with mixing depth, indicating that mixing depth is one of the important driving factors influencing water environment. According to mixing depth, a year can be divided into three different periods, including the thermally stratified period, isothermally mixed period, and transition period between them. When considering the former two different periods separately, mixing depth had no correlation with the phytoplankton biovolume. However, over the whole year a significant correlation was observed, which indicated that the influence of mixing depth on phytoplankton growth in the Zeya reservoir still followed Diehl’s theory. Furthermore, according to the steady-state assumption, a unimodal curve (mixing depth—phytoplankton biovolume) with a significant peak appearing at a mixing depth of 2 m was observed, closely agreeing with Diehl’ prediction.     

How do sinking phytoplankton species manage to persist?

Phytoplankton require light for photosynthesis. Yet, most phytoplankton species are heavier than water and therefore sink. How can these sinking species persist? Somehow, the answer should lie in the turbulent motion that redisperses sinking phytoplankton over the vertical water column. Here, we show, using a reaction-advection-diffusion equation of light-limited phytoplankton, that there is a turbulence window sustaining sinking phytoplankton species in deep waters. If turbulent diffusion is too high, phytoplankton are mixed to great depths, and the depth-averaged light conditions are too low to allow net positive population growth. Conversely, if turbulent diffusion is too low, sinking phytoplankton populations end up at the ocean floor and succumb in the dark. At intermediate levels of turbulent diffusion, however, phytoplankton populations can outgrow both mixing rates and sinking rates. In this way, the reproducing population as a whole can maintain a position in the well-lit zone near the top of the water column, even if all individuals within the population have a tendency to sink. This theory unites earlier classic results by Sverdrup and Riley as well as our own recent findings and provides a new conceptual framework for the understanding of phytoplankton dynamics under the influence of mixing processes.     

Climate change and the timing, magnitude, and composition of the phytoplankton spring bloom

In this article, we show by mesocosm experiments that winter and spring warming will lead to substantial changes in the spring bloom of phytoplankton. The timing of the spring bloom shows only little response to warming as such, while light appears to play a more important role in its initiation. The daily light dose needed for the start of the phytoplankton spring bloom in our experiments agrees well with a recently published critical light intensity found in a field survey of the North Atlantic (around 1.3 mol photons m^?2 day^?1). Experimental temperature elevation had a strong effect on phytoplankton peak biomass (decreasing with temperature), mean cell size (decreasing with temperature) and on the share of microplankton diatoms (decreasing with temperature). All these changes will lead to poorer feeding conditions for copepod zooplankton and, thus, to a less efficient energy transfer from primary to fish production under a warmer climate.     

Phytoplankton depth profiles and their transitions near the critical sinking velocity

We consider a simple phytoplankton model introduced by Shigesada and Okubo which incorporates the sinking and self-shading effect of the phytoplankton. The amount of light the phytoplankton receives is assumed to be controlled by the density of the phytoplankton population above the given depth. We show the existence of non-homogeneous solutions for any water depth and study their profiles and stability. Depending on the sinking rate of the phytoplankton, light intensity and water depth, the plankton can concentrate either near the surface, at the bottom of the water column, or both, resulting in a “double-peak” profile. As the buoyancy passes a certain critical threshold, a sudden change in the phytoplankton profile occurs. We quantify this transition using asymptotic techniques. In all cases we show that the profile is locally stable. This generalizes the results of Shigesada and Okubo where infinite depth was considered.      

Phytoplankton development and turbulent mixing in Lake Kinneret (1992-1996)

We have utilized data from a recently developed three-dimensionalvelocity fluctuation meter to compute the dissipation of turbulentkinetic energies (TKE) and the intensity of turbulent mixingin horizontal and vertical planes in the pelagic, epilimnicwater of Lake Kinneret, Israel. These characteristics of wind-inducedturbulent movement have been monitored from January 1992 throughDecember 1996. The turbulence parameters were strongly correlatedto wind energy inputs, calculated daily as 5 day cumulativeinputs. There have been dramatic changes in the annual and seasonaldevelopment of phytoplankton, together with unusually high levelsof primary production in this lake since 1994. We observed differentpatterns of vertical and horizontal turbulent movement and ofTKE dissipation rates during the years when ‘unusual’phytoplankton development occurred (1994–1996) comparedto ‘normal’ years (1992, 1993). The first appearanceof the filamentous cyanobacterium Aphanizomenon in this lakein August–September 1994 coincided with a period of markedlylower rates of TKE dispersion and a shift from vertical to horizontaldominance of the turbulent eddy spins. The absence of a regularwinter-spring bloom of the dinoflagellate, Peridinium, in 1996occurred when dissipation rates of TKE were extremely high,while record high amounts of dinoflagellates (1994, 1995) appearedwhen dissipation rates were very low. Correlations were shownbetween phytoplankton parameters (chlorophyll, primary productionand the ratio of primary production to chlorophyll) and boththe dissipation rate of TKE and the intensity of water turbulentmixing in the vertical plane. We suggest that the changes inthe ‘turbulence climate’ of Lake Kinneret were animportant factor in determining shifts in phytoplankton successionand the population composition of the algal assemblage.     

Comparative analysis of nutrients,chlorophyll and transparency in two large shallow lakes (Lake Taihu,P.R. China and Lake Okeechobee,USA)

This article compares limnological attributes of two of the world’s largest shallow lakes—Lake Okeechobee in Florida, USA and Lake Taihu in P.R. China. Both the systems support an array of ecological and societal values including fish and wildlife habitat, public water supply, flood protection, and recreation. Both have extensive research programs, largely because of concern regarding the lakes’ frequent cyanobacterial blooms. By evaluating these systems together, we compare and contrast properties that can generally advance the understanding and management of large shallow lowland lakes. Because of shallow depth, long fetch, and unconsolidated mud sediments, water chemistry, and transparency in both the lakes are strongly influenced by resuspended sediments that affect light and nutrient conditions. In the central region of both the lakes, where depth is the greatest, evaluation of limiting factors by a trophic state index approach indicates that light most often limits phytoplankton biomass. In contrast, the more sheltered shoreline areas of both the lakes display evidence of nitrogen (N) limitation, which also has been confirmed in nutrient assays conducted in earlier studies. This N limitation most likely is a result of excessive levels of phosphorus (P) that have developed in the lakes due to high external loads over recent decades and the currently high internal P recycling. Comparisons of these lakes show that Lake Taihu has higher N than, similar total phosphorus (TP) and similar light conditions to that of Lake Okeechobee, but less chlorophyll a (CHL). The latter may be as a result of lower winter temperatures in Lake Taihu (around 5°C) compared to Lake Okeechobee (around 15°C), which could reduce phytoplankton growth and abundance through the other seasons of the year. In these systems, the important role of light, temperature, and nutrients in algal bloom dynamics must be considered, especially due to possible adverse and unintended effects that might occur with projects such as sediment removal, and in the long term, in regard to buffering lake responses to external load reduction. Handling editor: D. Hamilton     

Implications of seasonal mixing for phytoplankton production and bloom development

Based on a 1D model considering phytoplankton and nutrients in a vertical water column, we investigate the consequences of temporal and spatial variations in turbulent mixing for phytoplankton production and biomass. We show that in seasonally mixed systems, the processes controlling phytoplankton production and the sensitivity of phytoplankton abundance to ambient light, trophic state and mixed-layer depth differ substantially from those at steady state in systems with time-constant diffusivities. In seasonally mixed systems, the annually replenished nutrient pool in the euphotic zone is an important factor for phytoplankton production supporting bloom development, whereas without winter mixing, production mainly depends on the diffusive nutrient flux during stratified conditions. Seasonal changes in water column production are predominantly determined by seasonal changes in phytoplankton abundance, but also by seasonal changes in specific production resulting from the transport of nutrients, the exploitation of the nutrient pool and the increase in light shading associated with phytoplankton growth. The interplay between seasonal mixing and the vertical distribution of mixing intensities is a key factor determining the relative importance of the processes controlling phytoplankton production and the sensitivity of the size and timing of the annual maximum phytoplankton abundance to the abiotic conditions.     

A model of phytoplankton blooms

A simple model that describes the dynamics of nutrient-driven phytoplankton blooms is presented. Apart from complicated simulation studies, very few models reported in the literature have taken this "bottom-up" approach. Yet, as discussed and justified from a theoretical standpoint, many blooms are strongly controlled by nutrients rather than by higher trophic levels. The analysis identifies an important threshold effect: a bloom will only be triggered when nutrients exceed a certain defined level. This threshold effect should be generic to both natural blooms and most simulation models. Furthermore, predictions are given as to how the peak of the bloom Pmax is determined by initial conditions. A number of counterintuitive results are found. In particular, it is shown that increasing initial nutrient or phytoplankton levels can act to decrease Pmax. Correct predictions require an understanding of such factors as the timing of the bloom and the period of nutrient buildup before the bloom.     

Spring bloom dynamics in Kongsfjorden,Svalbard: nutrients,phytoplankton, protozoans and primary production

The marine ecosystem in Kongsfjorden (79°N), a glacial fjord in Svalbard, is to a large extent well known with regard to hydrography, mesozooplankton and higher trophic levels. Research on primary production and lower trophic levels is still scare and especially investigations from winter and spring periods. The spring bloom dynamics in Kongsfjorden were investigated in 2002. The development in nutrient conditions, phytoplankton, protozoans and primary production were followed from 15 April until 22 May. The winter/spring in 2002 was categorized as a cold year with sea ice cover and water masses dominated by local winter-cooled water. The spring bloom started around 18 April and lasted until the middle of May. The bloom probably peaked in late April, but break-up of sea ice made it impossible to sample frequently in this period. Diatoms dominated the phytoplankton assemblage. We estimated the total primary production during the spring bloom in 2002 to range 27–35 g C m⁻². There was a mismatch situation between the mesozooplankton and the phytoplankton spring bloom in 2002.     

Phytoplankton competition in deep biomass maximum

Resource competition in heterogeneous environments is still an unresolved problem of theoretical ecology. In this article, I analyze competition between two phytoplankton species in a deep water column, where the distributions of main resources (light and a limiting nutrient) have opposing gradients and co-limitation by both resources causes a deep biomass maximum. Assuming that the species have a trade-off in resource requirements and the water column is weakly mixed, I apply the invasion threshold analysis (Ryabov and Blasius, Ecol Lett 14:220–228, 2011) to determine relations between environmental conditions and phytoplankton composition. Although species deplete resources in the interior of the water column, the resource levels at the bottom and surface remain high. As a result, the slope of resources gradients becomes a new crucial factor which, rather than the local resource values, determines the outcome of competition. The value of resource gradients nonlinearly depend on the density of consumers. This leads to complex relationships between environmental parameters and species composition. In particular, it is shown that an increase of both the incident light intensity or bottom nutrient concentrations favors the best light competitors, while an increase of the turbulent mixing or background turbidity favors the best nutrient competitors. These results might be important for prediction of species composition in deep ocean.     

A five-year study of autotrophic winter picoplankton in Lake Balaton,Hungary

Picoeukaryotes dominate the phytoplankton of Lake Balaton—the largest shallow lake in Central Europe—in the winter period. We examined the annual dynamics of picoplankton abundance and composition in the lake in order to establish if the picoeukaryotes merely survive the harsher winter conditions or they are able to grow in the ice-covered lake when the entire phytoplankton is limited by low light and temperature. Lake Balaton has an annual temperature range of 1–29°C, and it is usually frozen between December and February for 30–60 days. In the spring-autumn period phycocyanin and phycoerythrin rich Cyanobacteria are the dominant picoplankters, and picoeukaryotes are negligible. Our five-year study shows the presence of three types of picophytoplankton assemblages in Lake Balaton: (1) Phycoerythrin-rich Cyanobacteria—the dominant summer picoplankters in the mesotrophic lake area; (2) Phycocyanin-rich Cyanobacteria—the most abundant summer picoplankters in the eutrophic lake area and; (3) Picoeukaryotes—the dominant winter picoplankters in the whole lake. The observed winter abundance of picoeukaryotes was high (up to 3 × 10⁵ cells ml⁻¹), their highest biomass (520 μg l⁻¹) exceeded the maximum summer biomass of picocyanobacteria (500 μg l⁻¹). Our results indicate that the winter predominance of picoeukaryotes is a regular phenomenon in Lake Balaton, irrespective of the absence or presence of the ice cover. Picoeukaryotes are able to grow at as low as 1–2°C water temperature, while the total phytoplankton biomass show the lowest annual values in the winter period. In agreement with earlier findings, the contribution of picocyanobacteria to the total phytoplankton biomass in Lake Balaton is inversely related to the total phytoplankton biomass, whereas no such relationship was observable in the case of picoeukaryotes.     

Observations on the relation between phytoplankton diversity and environmental factors in the Vaal River at Balkfontein,South Africa

The relationship between specific environmental factors as independent variables and temporal changes in phytoplankton community structure in the Vaal River (a turbid system) during 1984 was investigated by employing different diversity indices. Temporal changes in community structure reflected temporal changes in certain environmental factors. Phytoplankton diversity, measured with Shannon-Wie H' and Hurlbert PIE indices, was related firstly to discharge and discharge derived variables (such as SO₄, Si, N and P loading) and secondly to turbidity derived variables (such as euphotic zone depth). Discharge appears to be of prime importance in affecting diversity. Observations were made that shed new light on conditions contributing to the development of an August peak (dominated by Stephanodiscus hantzschii fo. tenuis and Micractinium pusillum) in phytoplankton concentration. Increased environmental stress may reduce the number of sensitive species, thus reducing interspecific competition between tolerant species which could then exploit the — for them — more favourable conditions resulting in an increase in their numbers to peak concentrations.     

Modelling phytoplankton productivity in turbid waters with small euphotic to mixing depth ratios

A model which was developed and calibrated for predicting algalgrowth in mass cultures, was modified for natural systems. Inturbid systems the ratio of euphotic to aphotic depth is usuallysmall and mixing may exceed the compensation depth. In orderto compensate for the various light regimes to which the phytoplanktonwould be subjected, the losses due to dark respiration weremodified so that the effective light history would determinethe actual rate. The efficiency of light utilization also changesunder different light regimes and the model was modified totake these variations into account. Both these modificationsresulted in different production profiles being generated forthe same surface conditions, but with different mixing depths,where the phytoplankton become more efficient as the light regimedeteriorates (i.e. less respiration and greater light utilizationefficiency). A further consequence is that the ‘criticalmixing depth’ is {small tilde}2.5 times greater than thatwhich was previously accepted, being {small tilde}20 times theeuphotic depth. The model predicted productivities to within90% of observed rates. It was also suggested how the predictionscould be used to determine the extent of nutrient limitation.The predictions also have biomanipulatory consequences, as alterationsof the light regime through the addition of non-photosynthesizingmaterials, under certain conditions, may even result in a stimulationof phytoplankton productivity.     

Spring bloom succession,grazing impact and herbivore selectivity of ciliate communities in response to winter warming

This study aimed at simulating different degrees of winter warming and at assessing its potential effects on ciliate succession and grazing-related patterns. By using indoor mesocosms filled with unfiltered water from Kiel Bight, natural light and four different temperature regimes, phytoplankton spring blooms were induced and the thermal responses of ciliates were quantified. Two distinct ciliate assemblages, a pre-spring and a spring bloom assemblage, could be detected, while their formation was strongly temperature-dependent. Both assemblages were dominated by Strobilidiids; the pre-spring bloom phase was dominated by the small Strobilidiids Lohmaniella oviformis, and the spring bloom was mainly dominated by large Strobilidiids of the genus Strobilidium. The numerical response of ciliates to increasing food concentrations showed a strong acceleration by temperature. Grazing rates of ciliates and copepods were low during the pre-spring bloom period and high during the bloom ranging from 0.06 (Δ0°C) to 0.23 day⁻¹ (Δ4°C) for ciliates and 0.09 (Δ0°C) to 1.62 day⁻¹ (Δ4°C) for copepods. During the spring bloom ciliates and copepods showed a strong dietary overlap characterized by a wide food spectrum consisting mainly of Chrysochromulina sp., diatom chains and large, single-celled diatoms. Priority programme of the German Research Foundation—contribution 4.     

Phytoplankton response to a changing climate

Phytoplankton are at the base of aquatic food webs and of global importance for ecosystem functioning and services. The dynamics of these photosynthetic cells are linked to annual fluctuations of temperature, water column mixing, resource availability, and consumption. Climate can modify these environmental factors and alter phytoplankton structure, seasonal dynamics, and taxonomic composition. Here, we review mechanistic links between climate alterations and factors limiting primary production, and highlight studies where climate change has had a clear impact on phytoplankton processes. Climate affects phytoplankton both directly through physiology and indirectly by changing water column stratification and resource availability, mainly nutrients and light, or intensified grazing by heterotrophs. These modifications affect various phytoplankton processes, and a widespread advance in phytoplankton spring bloom timing and changing bloom magnitudes have both been observed. Climate warming also affects phytoplankton species composition and size structure, and favors species traits best adapted to changing conditions associated with climate change. Shifts in phytoplankton can have far-reaching consequences for ecosystem structure and functioning. An improved understanding of the mechanistic links between climate and phytoplankton dynamics is important for predicting climate change impacts on aquatic ecosystems.     

Sedimentation loss of phytoplankton cells from the mixed layer: effects of turbulence levels

In this paper, we describe a study of the role of turbulencein the loss by sedimentation of phytoplankton cells from themixed layer. The approach presented allows the quantificationof the sedimentation rate of phytoplankton in the whole rangeof turbulence levels of this layer. Two types of phytoplanktercan be distinguished according to the effect that turbulencecan exert on their sedimentation rate. The rate of those cellswhose settling velocity is lower than –1 m day^–1will not be modified by turbulence. The sedimentation rate ofcells with higher settling velocities can, however, be modifiedby the level of turbulence. A set of dimensionless numbers isgiven to delimit several processes that are important in thedynamics of phytoplankton sedimentation in a turbulent regime.The use of these dimensionless numbers suggests that an increasein the turbulence level in the mixed layer does not always implya decrease in the sedimentation rate of phytoplankton cells.