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1.
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.  相似文献   

2.
3.
Kiørboe T 《Oecologia》2008,155(1):179-192
The performance of individual phytoplankton species is strongly governed by the thermal stratification’s impact on vertical mixing within the water column, which alters the position of phytoplankton relative to nutrients and light. The present study documents shifts in phytoplankton structure and vertical positioning that have accompanied intensified long-term stratification in a natural ecosystem. Ordination analysis is used to extract gradients in phytoplankton composition in Lake Tahoe, an extremely nutrient-poor lake, over a 23-year period of records. Community structure in the 1980s was associated most strongly with resource availability (low nitrogen to phosphorus ratios, deeper euphotic zone depth), while intensified stratification dominated the phytoplankton structure since the late 1990s. Within diatoms, small-sized cells increased with reduced mixing, suggesting that suppressed turbulence provides them with a competitive advantage over large-sized cells. Among the morphologically diverse chlorophytes, filamentous and coenobial forms were favored under intensified stratification. The selection for small-sized diatoms is accompanied by a shoaling trend in their vertical position in the water column. In contrast, the motile flagellates displayed a deeper vertical positioning in recent years, indicating that optimal growth conditions shifted likely due to reduced upwelling of nutrients. As the thermal stratification of lakes and oceans is strongly linked to climate variables, the present study confirms that climate warming will alter phytoplankton structure and dynamics largely through effects on nutrient availability and sinking velocities. Intensified stratification should favor the expansion of small-sized species and species with the capability of buoyancy regulation, which may alter primary productivity, nutrient recycling, and higher trophic productivity. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.  相似文献   

4.
Water-column mixing is known to have a decisive impact on plankton communities. The underlying mechanisms depend on the size and depth of the water body, nutrient status and the plankton community structure, and they are well understood for shallow polymictic and deep stratified lakes. Two consecutive mixing events of similar intensity under different levels of herbivory were performed in enclosures in a shallow, but periodically stratified, eutrophic lake, in order to investigate the effects of water-column mixing on bacteria abundance, phytoplankton abundance and diversity, and rotifer abundance and fecundity. When herbivory by filter-feeding zooplankton was low, water-column mixing that provoked a substantial nutrient input into the euphotic zone led to a strong net increase of bacteria and phytoplankton biomass. Phytoplankton diversity was lower in the mixed enclosures than in the undisturbed ones because of the greater contribution of a few fast-growing species. After the second mixing event, at a high biomass of filter-feeding crustaceans, the increase of phytoplankton biomass was lower than after the first mixing, and diversity remained unchanged because enhanced growth of small fast-growing phytoplankton was prevented by zooplankton grazing. Bacterial abundance did not increase after the second mixing, when cladoceran biomass was high. Changes in rotifer fecundity indicated a transmission of the phytoplankton response to the next trophic level. Our results suggest that water-column mixing in shallow eutrophic lakes with periodic stratification has a strong effect on the plankton community via enhanced nutrient availability rather than resuspension or reduced light availability. This fuels the basis of the classic and microbial food chain via enhanced phytoplankton and bacterial growth, but the effects on biomass may be damped by high levels of herbivory. Received: 3 May 1999 / Accepted: 13 April 2000  相似文献   

5.
SUMMARY. I. Movement of 33P from hypolimnion to epilimnion in a small, dystrophic lake was investigated using small-diameter experimental tubes enclosing thermally stratified water columns. This approach was made possible by the extremely sharp stratification found in such lakes, in which the euphotic zone closely coincides with the epilimnion.
2. The vertical distribution of inorganic phosphorus in the lake showed a sharp increase across the thermocline so that enhanced concentrations were available to phytoplankton just below the thermocline. Inorganic nitrogen concentrations did not show such a marked relation to thermal stratification.
3. One abundant motile alga ( Cryptomonas marssonii ) showed striking and regular vertical migrations in the lake, moving below the thermocline at night and returning to the surface waters in early morning. These migrations took cells across a 10°C temperature gradient. Non-motile phytoplankton showed constant vertical distributions.
4. In the experimental tubes an upward movement of phosphorus took place from hypolimnion to epilimnion which was only attributable to transport by phytoplankton cells undertaking active vertical migrations. No equivalent movement of phosphorus occurred in control tubes from which algae were absent.
5. The possible significance of such nutrient retrieval is discussed with reference to plankton phosphorus budgets and competition between phytoplankton species.  相似文献   

6.
7.
Lake Baikal, Russian Siberia, was sampled in July 1990 during the period of spring mixing and initiation of thermal stratification. Vertical profiles of temperature, dissolved nutrients (nitrate and soluble reactive phosphorus), phytoplankton biomass, and primary productivity were determined in an eleven-station transect encompassing the entire 636 km length of the lake. Pronounced horizontal variability in hydrodynamic conditions was observed, with the southern region of the lake being strongly thermally stratified while the middle and north basins were largely isothermal through July. The extent of depletion of surface water nutrients, and the magnitude of phytoplankton biomass and productivity, were found to be strongly correlated with the degree of thermal stratification. Horizontal differences likely reflected the contribution of two important factors: variation in the timing of ice-out in different parts of the lake (driving large-scale patterns of thermal stratification and other limnological properties) and localized effects of river inflows that may contribute to the preliminary stabilization of the water column in the face of intense turbulent spring mixing (driving meso-scale patterns). Examination of the relationships between surface water inorganic N and P depletion suggested that during the spring and early summer, phytoplankton growth in unstratified portions of the lake was largely unconstrained by nutrient supplies. As summer progressed, the importance of co-limitation by both N and P became more apparent. Uptake and regeneration rates, measured directly using the stable isotope 15N, revealed that phytoplankton in stratified portions of the lake relied primarily on NH4 as their N source. Rates of NH4 regeneration were in approximate equilibrium with uptake; both processes were dominated by organisms <2 µm. This pattern is similar to that observed for oligotrophic marine systems. Our study underscores the importance of hydrodynamic conditions in influencing patterns of biological productivity and nutrient dynamics that occur in Lake Baikal during its brief growing season.  相似文献   

8.
A knowledge of diel variation and the vertical distribution of phytoplankton communities may contribute to a better understanding of the driving factors of key species. Applying functional-group classification provides important information on the causes of species selection in the pelagic community. The diel variation of phytoplankton functional groups was analysed during an autumnal stratification period with the aim of understanding their changes in the vertical position related to light, mixing regime and grazing pressure. Phytoplankton and zooplankton communities were sampled every 4 h during a 24-h period in a vertical profile in a subtropical meso-eutrophic reservoir. Strong stratification during a 24-h cycle and a mixed clear epilimnion with partial atelomixis marked the autumn season in the Faxinal reservoir, southern Brazil. The highest phytoplankton densities and biomass were found during the second part of the day, a general pattern reported in the literature, and may be explained by zooplankton dynamics. During the 24-h cycle, phytoplankton functional groups lacking a self-regulating capacity and those able to regulate their vertical position were vertically segregated in the lake. The diel behaviour of both groups was driven by the mixing regime (including atelomixis), light and zooplankton grazing pressure.  相似文献   

9.
The vertical distribution of phytoplankton in stratified water columns   总被引:2,自引:0,他引:2  
What determines the vertical distribution of phytoplankton in different aquatic environments remains an open question. To address this question, we develop a model to explore how phytoplankton respond through growth and movement to opposing resource gradients and different mixing conditions. We assume stratification creates a well-mixed surface layer on top of a poorly mixed deep layer and nutrients are supplied from multiple depth-dependent sources. Intraspecific competition leads to a unique strategic equilibrium for phytoplankton, which allows us to classify the distinct vertical distributions that can exist. Biomass can occur as a benthic layer (BL), a deep chlorophyll maximum (DCM), or in the mixed layer (ML), or as a combination of BL+ML or DCM+ML. The ML biomass can be limited by nutrients, light, or both. We predict how the vertical distribution, relative resource limitation, and biomass of phytoplankton will change across environmental gradients. We parameterized our model to represent potentially light and phosphorus limited freshwater lakes, but the model is applicable to a broad range of vertically stratified systems. Increasing nutrient input from the sediments or to the mixed layer increases light limitation, shifts phytoplankton towards the surface, and increases total biomass. Increasing background light attenuation increases light limitation, shifts the phytoplankton towards the surface, and generally decreases total biomass. Increasing mixed layer depth increases, decreases, or has no effect on light limitation and total biomass. Our model is able to replicate the diverse vertical distributions observed in nature and explain what underlying mechanisms drive these distributions.  相似文献   

10.
Phytoplankton of the Barents Sea - the end of a growth season   总被引:2,自引:2,他引:0  
 Few phytoplankton investigations have been carried out at the end of the growth season, particularly in the Arctic. In the present study, we monitored the phytoplankton distribution in relation to environmental conditions in the Barents Sea in September 1988 and October 1987. An ice-edge bloom was found in September at 80° N in a stratified meltwater layer, lasting until new ice formation and southward advection of the ice cover commenced in the middle of the month. Phytoplankton populations in the marginal ice zone at this time were not nutrient limited, but biomass was probably reduced due to grazing by small copepods. Lower chl/C and chl/N ratios in the phytoplankton above the pycnocline than below in September indicated light-adapted populations. In October the particulate matter was rich in carbon, but had low chlorophyll content, indicating high levels of detritus. The hydrographic conditions in October differed greatly from those observed in September. The combination of freezing and mixing resulted in higher salinity and nutrient concentrations, and caused a homogeneous distribution, as well as reduction, of the phytoplankton stocks in the upper water column. During late October, low incoming radiation, combined with deep vertical mixing, resulted in light-limiting conditions for the algae, eventually stopping photosynthesis and terminating the growth season in the northern Barents Sea. Received: 1 March 1996/Accepted: 19 May 1996  相似文献   

11.
Nixdorf  Brigitte 《Hydrobiologia》1994,(1):173-186
The polymictic properties of Lake Müggelsee, a eutrophic shallow lake in Berlin, are described by the water column stability (N 2) and gradients in saturation of oxygen at the deepest site of the lake (7.5 m). Mixing and stratification changed irregularly up to 7 times during the vegetation season (April to September), as was indicated by all of the stratification parameters. Thermally stable conditions generally lasted 1–2 weeks. A maximum of 5 weeks stratification was observed in 1982.In order to investigate the response of algal development, the internal rates of change of the dominant algal species in the lake during the vegetation period were estimated from weekly measurements of phytoplankton biomass from 1980 to 1990. The necessity taking a mixed sample in a shallow lake is discussed. The polymictic properties favoured the development of specific blue-green algal species; there dominance was also favoured by the trophic conditions. Among the dominant blue-greens the growth of Limnothrix redekei was independent of polymixis whereas stratification supported the starting conditions for the summer blue-greens Aphanizomenon flos-aquae and Planktothrix agardhii. After these algae reached a distinct level of biomass, they grew under mixing as well as under stratified conditions.For the development of solitary centric diatoms during summer regulation by growth restriction through nutrient limitation, esp. dissolved silicon was more important. However, Melosira sp. developed well under stratified conditions but collapsed due to increased sinking losses when the water column became too stable.An attempt is made to apply Reynolds' possibility matrix of the most likely phytoplankton assemblages as a function of nutrients and mixing in the shallow Lake Müggelsee.  相似文献   

12.
ANTARCTIC AQUATIC ECOSYSTEMS AS HABITATS FOR PHYTOPLANKTON   总被引:10,自引:0,他引:10  
1. The Southern Ocean is a large-scale, relatively homogeneous upwelling ecosystem whose phytoplankton apparently grows suboptimally over much of its area. By contrast there is a wide variety of freshwater habitats in the Antarctic and in some of these phytoplankton growth efficiency is very high. The two habitats share similar temperature and irradiance regimes, but differ markedly in availability of inorganic nutrients, in grazing pressure and in the time- and space-scales on which various physical processes act. 2. Concentrations of inorganic nutrients in the marine ecosystem have been represented as being in excess of phytoplankton requirements, but the ionic composition of some nutrient pools may not conform to phytoplankton preferences. 3. Nutrient-limitation determines phytoplankton production in Antarctic lakes and gives rise to gross differences between lakes. 4. Irradiance in the water column varies greatly over the year in both marine and freshwater ecosystems. Most algae are shade-adapted, with the ability to utilize low irradiance but with sub-optimal response to high irradiance. However, local phytoplankton maxima may attain very high carbon fixation and growth rates. 5. Consistently low temperatures characterize both systems. Their effects on photo-synthetic carbon uptake mirror shade-adaptation. Division rates of marine phytoplankton may however be very much higher than predicted for ambient temperatures. 6. Vertical mixing is important in both ecosystems and influences the environment experienced by phytoplankton cells. This appears to have little effect on the average performance of phytoplankton in the strongly mixed surface water column of the Southern Ocean, where the mixed depth may exceed 100 m. This can be related partly to the shade-adapted photosynthetic response. Euphotic depths range from 20 to 100 m. 7. Strong vertical mixing under ice-free conditions in lakes may maximize photosynthetic efficiency, whilst distinct vertical stratification in permanently ice-covered lakes gives rise to segregation of nutrient uptake and regeneration. 8. Physical removal of phytoplankton biomass by grazing is locally important in the Southern Ocean, in contrast to the estimated mean mesoscale impact of grazing. Vertical sedimentation losses appear important in the context of mixing depth and generation time, and may be modified by vertical circulation of water. 9. Loss of phytoplankton biomass from lakes during the ice-free period is dominated by physical removal via the lake outflow. Grazing is generally unimportant, except where larvae of otherwise nektobenthic zooplankton hatch in synchrony with a phytoplankton maximum. Sedimentation is important under ice-cover.  相似文献   

13.
The present study was carried out in Faxinal Reservoir, a warm monomictic, meso-eutrophic reservoir in subtropical southern Brazil, with a long-standing, well-stratified condition, low epilimnetic nutrient concentrations, and a relatively clear epilimnion. In this study, we analyzed the dynamics of the phytoplankton functional groups, recognizing their driving forces in Faxinal Reservoir. Samples were taken at monthly intervals from January 2004 to January 2005 in surface waters. According to the reservoir’s mixing regime, three periods were identified during the study: stratification 1 (January–May 2004); mixing period (June–August 2004); and stratification 2 (September 2004–January 2005). The nutrient dynamics were driven by the mixing regime. The H1, F, and C phytoplankton functional groups were the most important in biomass, mainly represented by the N-fixing cyanobacterium Anabaena crassa, the colonial green alga with thick mucilaginous sheaths Nephrocytium sp., and the diatom Asterionella formosa, respectively. Tendencies pointed out by redundancy analysis (RDA) indicated that the mixing regime was the main determining factor of the seasonal dynamics of the phytoplankton community. The dominant functional groups showed a close relationship with the relative water-column stability (RWCS), and also, as a consequence of the mixing regime, with nutrient availability. The study also revealed the important role of physical processes in the seasonal gradient, in selecting for phytoplankton functional groups and, consequently, in the assessment of ecological status. Q index (assemblage index) of water quality based on functional groups revealed ecological status varying from very poor to tolerable in the stratification 1 period and from tolerable to medium in the mixing and stratification 2 periods. Handling editor: Judit Padisak  相似文献   

14.
A model of phytoplankton dynamics within a water column was analyzed with special consideration on the existence of a bifurcation set in the parameter space. We considered two resources, light and a limiting nutrient, for phytoplankton growth and assumed that the water column is separated into two layers by thermal and/or density stratification. It was shown that there exists a bifurcation set in the parameter space when the growth function meets several conditions that are general for growth functions of two essential resources. Specifically, these conditions include that a less abundant of the two resources limits the growth while the effect of the other is sufficiently small. Folded structure with two stable states separated by one unstable state appears in the catastrophe manifold when parameters move to a certain direction with a certain curvature from a point in the bifurcation set. These results suggest that occurrence of discontinuous transition between two alternative vertical patterns is possible nature of phytoplankton dynamics within a stratified water column.  相似文献   

15.
1. The impact of long thermal stratification events on some key properties in a polymictic lake was studied by determining the mixing regime of Müggelsee, Germany, using water temperature profiles taken hourly over 4 years. The period included two exceptional summer heatwaves. 2. Long thermal stratification events lasted from about 1 week to 2 months, and exhibited a high variability in thermocline depth and stratification intensity within and between events. 3. During stratification events, hypolimnetic oxygen concentrations strongly decreased while hypolimnetic SRP accumulation increased, depending on the duration and intensity of stratification and on hypolimnetic water temperature. 4. The impact of stratification on the functional phytoplankton composition increased with increasing stratification duration, but was rather different for the heatwaves. 5. Stratification events were followed by strong nutrient pulses into the euphotic zone and intense phytoplankton growth, particularly after the heatwaves. Hence, the influence of the climate extremes counteracted effects of reduced external nutrient loading.  相似文献   

16.
1. Wind is considered the dominant factor controlling phytoplankton distribution in lentic environments. In canyon‐shaped reservoirs, wind tends to blow along the main axis generating internal seiches and advective water movements that jointly with biological features of algae can produce a heterogeneous phytoplankton distribution. Turbulence generated by wind stress and convection will also affect the vertical distribution of algae, depending on their sinking properties. 2. We investigated the vertical and horizontal distribution of phytoplankton during the stratification period in Sau Reservoir (NE Spain). Sites along the main reservoir axis were sampled every 4 h for 3 days, and profiles of chlorophyll‐a and temperature were made using a fluorescent FluoroProbe, which can discriminate among the main algal groups. Convective and wind shear velocity scales, and energy dissipation were calculated from meteorological data, and simulation experiments were performed to describe non‐measured processes, like vertical advection and sinking velocity of phytoplankton. 3. Wind direction changed from day to night, producing a diel thermocline oscillation and an internal seiche. Energy dissipation was moderate during the night, and mainly attributed to convective cooling. During the day the energy dissipation was entirely attributed to wind shear, but values indicated low turbulence intensity. 4. The epilimnetic algal community was mainly composed of diatoms and chlorophytes. Chlorophytes showed a homogeneous distribution on the horizontal and vertical planes. Diatom horizontal pattern was also homogeneous, because the horizontal advective velocities generated by wind forcing were not high enough to develop phytoplankton gradients along the reservoir. 5. Diatom vertical distribution was heterogeneous in space and time. Different processes dominated in different regions of the reservoir, due to the interaction between seiching and the daily cycle of convective‐mediated turbulence. As the meteorological forcing followed a clear daily pattern, we found very different diatom sedimentation dynamics between day and night. Remarkably, these dynamics were asynchronous in the extremes of the seiche, implying that under the same meteorological forcing a diatom population can show contrasting sedimentation dynamics at small spatial scales (approximately 103 m). This finding should be taken into account when interpreting paleolimnological records from different locations in a lake. 6. Vertical distribution of non‐motile algae is a complex process including turbulence, vertical and horizontal advection, variations in the depth of the mixing layer and the intrinsic sinking properties of the organisms. Thus, simplistic interpretations considering only one of these factors should be regarded with caution. The results of this work also suggest that diatoms can persist in stratified water because of a synergistic effect between seiching and convective turbulence.  相似文献   

17.
1. The effects of wind events on phytoplankton dynamics were investigated in two temperate reservoirs. 2. Meteorological forcing, change in physical and chemical structure of the water column and biological responses of phytoplankton communities were followed for 3 weeks in three seasons. 3. Depending on the season, the phytoplankton response differed in response to nutrient and light conditions, and to the intensity of stratification and mixing. 4. We demonstrated that, on a time scale of a few days, wind events can modify phytoplankton dynamics, in terms of size structure and exported biomass. An increase of mixing favoured the largest size class and disadvantaged the smallest size class, while an increase in stratification had the opposite effects. The short‐term change in size structure was reflected in the sedimentary fluxes but with a time lag.  相似文献   

18.
We showed a catastrophic transition between a surface maximum pattern and a subsurface maximum pattern of phytoplankton in a water column by a mathematical model considering the vertical distribution of phytoplankton and two resources, light and a limiting nutrient. In our model, we assumed that a water column consists of two layers: a complete mixing layer above a seasonal thermocline and an incomplete mixing layer below it. From numerical calculation of the model, we obtained that there are two stable vertical patterns of phytoplankton over a certain range of parameters of the model: a pattern having its maximum below the thermocline and another having its maximum above the thermocline. As other models having multiple stable equilibria, our model also exhibits a hysteresis effect and catastrophic transition when one of the parameters of the model changes continuously. These results indicate the possibility of the existence of alternative equilibria of vertical patterns of phytoplankton even if the trophic status and physical condition of the water column are similar. Moreover, the catastrophic transition between the steady states suggests one of the possible mechanisms of autumn algal blooms.  相似文献   

19.
A one-dimensional numerical model and a two-dimensional numerical model of the hydrodynamic and thermal structure of Lake Shira during summer have been developed, with several original physical and numerical features. These models are well suited to simulate the formation and dynamics of vertical stratification and provide a basis for an ecological water-quality model of the lake. They allow for the quantification of the vertical mixing processes that govern not only the thermal structure but also the nutrient exchange, and more generally, the exchange of dissolved and particulate matter between different parts of the lake. The outcome of the calculations has been compared with the field data on vertical temperature and salinity distributions in Lake Shira. Lake Shira is meromictic and exhibits very stable annual stratification. The stratification is so stable because of the high salinity of the water. If the water in Lake Shira were fresh and other parameters (depth, volume, and meteorology) were the same, as now, the lake would be mixed in autumn. Using the newly developed models and using common meteorological parameters, we conclude that Lake Shira will remain stratified in autumn as long as the average salinity is higher than 3‰.  相似文献   

20.
Scale and frequency of changes in a lake’s physical structure, light dynamics, and availability of nutrients are closely related to phytoplankton ecology. Since phytoplankton assemblages were first described, phytoplankton ecologists concluded that these assemblages provide insight into phytoplankton responses to environmental changes. Objectives of this study were to investigate ecology of phytoplankton during a complete hydrological cycle in the deepest natural lake in Brazil, Dom Helvécio, and to sort species into the list of assemblages, checking its accordance with environmental changes in a tropical system within the middle Rio Doce Lake district, South-East Brazil. Canonical Correspondence Analysis, t-test, Mann–Whitney U-test, and Kruskal–Wallis test were used to analyze climatological, environmental, and plankton data, which were obtained monthly in 2002. A new phytoplankton assemblage, NA (atelomixis-dependent desmids), is suggested because atelomixis (robust movement of water occurring once a day) contributed to replacement of species in Dom Helvécio Lake. Stability of stratification, water chemistry, and composition of phytoplankton assemblages characterized two periods. The first period occurred in six rainy months (Jan–Mar and Oct–Dec) when the lake was stratified and phytoplankton was dominated by two assemblages: NA and F. The second period occurred in six dry months (Apr–Sep) when the lake was nonstratified and phytoplankton was dominated by four assemblages: S2, X1, A, and LO. Results suggest that phytoplankton in Dom Helvécio Lake was shaped by seasonal and daily changes of water temperature, even with its lower amplitude of variation within 2002 (El Niño year). These changes promoted water column stratification or mixing, reduced light, and increased nutrient availability. Temperature, therefore, is similarly important to phytoplankton ecology in tropical regions as it is in temperate ones. Sorting phytoplankton species into assemblages matched well with environmental changes and periods identified so it is also suggested that this can be further used as an appropriate tool to manage water quality when evaluating tropical lakes.  相似文献   

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