Warming promotes cold-adapted phytoplankton in temperate lakes and opens a loophole for Oscillatoriales in spring

The effects of the recent warming trend in many northern temperate lakes on the species composition of spring phytoplankton remain poorly understood, especially because a recent change in nutrients has complicated efforts, and previous studies have defined spring according to the calendar. We analysed data from 1979 to 2004 from Lake Müggelsee (Berlin, Germany), using physical and biological parameters to define the spring period. We show that a change in timing of spring plankton events in warm years led to the paradox of lower mean water temperatures during the growth period, favouring cold-adapted diatoms over cyanobacteria, and within the diatoms, some cold-adapted centric forms over pennate forms. Under high P : Si ratios, the increased time between phytoplankton and cladoceran peaks opened a loophole for filamentous cyanobacteria (Oscillatoriales) in warm years to establish dominance after the diatoms, which are silicate limited. Therefore, the warming trend promotes filamentous cyanobacteria, a well-known nuisance in eutrophic lakes, and surprisingly, cold-adapted diatoms.     

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.     

Climate change and the phytoplankton spring bloom: warming and overwintering zooplankton have similar effects on phytoplankton

Indoor mesocosms were used to study the combined effect of warming and of different densities of overwintering mesozooplankton (mainly copepods) on the spring development of phytoplankton in shallow, coastal waters. Similar to previous studies, warming accelerated the spring phytoplankton peak by ca. 1 day °C^?1 whereas zooplankton did not significantly influence timing. Phytoplankton biomass during the experimental period decreased with warming and with higher densities of overwintering zooplankton. Similarly, average cell size and average effective particle size (here: colony size) decreased both with zooplankton density and warming. A decrease in phytoplankton particle size is generally considered at typical footprint of copepod grazing. We conclude that warming induced changes in the magnitude and structure of the phytoplankton spring bloom cannot be understood without considering grazing by overwintering zooplankton.     

The phytoplankton of some gravel-pit lakes in Spain

The phytoplankton communities of thirteen adjacent gravel-pit lakes in the lower Jarama river watershed (Madrid, Spain), were studied during spring mixing and summer stratification.If different seasons, the phytoplankton responded to different environmental factors. During spring, the abundance of SRP (soluble reactive phosphorus) and existence of a certain thermal stability resulted in the development of a greater biomass in some lakes. During summer, however, excessively high temperatures adversely affected the communities of the warmer lakes. At the species level, the responses were diverse; ordination techniques enabled us to group them.Some similarities were observed in phytoplankton composition between lakes, possibly due to local dispersion between adjacent lakes (frequented by abundant waterfowl).     

Increasing importance of small phytoplankton in a warmer ocean

The macroecological relationships among marine phytoplankton total cell density, community size structure and temperature have lacked a theoretical explanation. The tiniest members of this planktonic group comprise cyanobacteria and eukaryotic algae smaller than 2 μm in diameter, collectively known as picophytoplankton. We combine here two ecological rules, the temperature–size relationship with the allometric size‐scaling of population abundance to explain a remarkably consistent pattern of increasing picophytoplankton biomass with temperature over the ?0.6 to 22 °C range in a merged dataset obtained in the eastern and western temperate North Atlantic Ocean across a diverse range of environmental conditions. Our results show that temperature alone was able to explain 73% of the variance in the relative contribution of small cells to total phytoplankton biomass regardless of differences in trophic status or inorganic nutrient loading. Our analysis predicts a gradual shift toward smaller primary producers in a warmer ocean. Because the fate of photosynthesized organic carbon largely depends on phytoplankton size, we anticipate future alterations in the functioning of oceanic ecosystems.     

Thermal stratification, nutrient dynamics, and phytoplankton productivity during the onset of spring phytoplankton growth in Lake Baikal, Russia

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.     

Further studies of a spring phytoplankton bloom in an enclosed experimental ecosystem

A detailed characterization is presented of the spring diatom bloom which occurred in the enclosed experimental ecosystem bags at Loch Ewe, Scotland, during March–April 1983. The nutrient condition and bacterial biomass of the water column, phytoplankton species distribution, gross biochemical composition and detailed lipid composition (lipid class, fatty acid and free sterol) of the phytoplankton are reported throughout the bloom period. The results are compared with results from previous years. The conclusions are that major changes take place in the biochemical composition of a rapidly growing diatom population which affect both the gross composition and also the more detailed lipid composition. Such changes can take place over a matter of days and appear to be very dependent upon available growth conditions. Both carbohydrate and lipids levels increase towards the end of the bloom as nitrate and silicate levels are depleted in the water. Neutral lipids are shown to be important lipid components of the phytoplankton populations and long-chain ω3 polyunsaturated fatty acids are found to be only minor components of the structural polar lipids. The fatty acid and sterol data are discussed in relation to present knowledge concerning phytoplankton lipid composition.     

Dynamics in bacterial surface properties of a natural bacterial community in the coastal North Sea during a spring phytoplankton bloom

The hydrophilic and hydrophobic properties of single cells of natural bacterioplankton communities were determined using a recently developed staining method combined with confocal laser scanning microscopy and advanced image analysis. On an average, about 50% of the bacterial cell area was covered by hydrophobic and only 16% by hydrophilic properties, while about 72% was covered by the genome. However, the size of these properties was independent of the bacterial cell size. Bacterial hydrophobicity was positively correlated with ambient NH(4)(+) concentrations and negatively correlated with overall bacterial abundance. The expression of hydrophilicity was more dynamic. Over the spring phytoplankton bloom, the bacterioplankton ratio(phil/phob) repeatedly reached highest values shortly before peaks in bacterioplankton abundance were observed, indicating a direct and fast response of bacterial surface properties, especially hydrophilicity, to changing environmental conditions. Compared to bacterial strains, recently studied with the same method, cells of marine bacterioplankton communities are much smaller and less frequently covered by hydrophobic or hydrophilic properties. While the percentage area covered by the genome is essentially the same, the percentage area covered by hydrophobic or hydrophilic properties is much smaller.     

Iron limits primary productivity during spring bloom development in the central North Atlantic

We present in situ biophysical measurements and bioassay experiments that demonstrate iron limitation of primary productivity during the spring bloom in the central North Atlantic. Mass balance calculations indicate that nitrate drawdown is iron (Fe)-limited and that aeolian Fe supply to this region cannot support maximal phytoplankton growth during the bloom. Using a simple simulation model, we show that relief of Fe limitation during the spring bloom can increase nitrate drawdown and, hence, new primary production, by 70%. We conclude that the episodic nature of iron supplied by dust deposition is an important factor controlling the dynamics of the spring bloom. From this, we hypothesize that variability in the timing and magnitude of the spring bloom in response to aeolian Fe supply will affect carbon drawdown and food web dynamics in the central North Atlantic.     

Effects of recent climate change on phytoplankton phenology in a temperate lake

1. A number of long-term studies have shown that spring biological events have advanced in recent decades and that this is a response to climate change. In lentic systems, changes in phytoplankton phenology have been attributed to various directly climate-related processes including changes in the onset and duration of thermal stratification, earlier ice-break up and increased water temperature. Both indirect climatic drivers and non-climate drivers such as elevated grazing pressure and nutrient enrichment can also affect phenology.
2. This study investigated whether phenological trends in phytoplankton could be detected in a relatively short time series in a shallow, ice-free, polymictic lake with a high annual discharge and whether any such trends could be causally explained.
3. It was found that the centre of gravity of the spring chlorophyll a bloom advanced significantly by 1.6 days per year over a 15-year period. This was accompanied by a significant increase in water temperature of 0.12 °C per year which is high compared to published rates of change over longer time series. No direct effects of ice cover, stratification or water discharge rates could be linked to the advancement of the spring bloom. Instead, the shift in timing was attributed to an advance in the timing of the dominant spring diatom, Aulacoseira spp., instigated by a temperature-driven increase in replication rate leading to an earlier onset of silica (SiO₂) limitation.     

Short-term variations in the physiological state of phytoplankton [2pt] in a shallow temperate estuary

Short-term changes in the photosynthetic carbon metabolism and physiological state of phytoplankton were studied over a summer fortnight-long period in the Urdaibai estuary (Bay of Biscay) and related to observed environmental patterns. Day-to-day variability in the hydrographical and biological features of the estuary during the study period was due to changes in meteorological and tidal conditions. Phytoplankton biomass and primary production increased with the improvement of weather, i.e., light conditions, during neap tides. Thus a mixed bloom of cryptophyceans, Euglena sp., and the dinoflagellate Peridinium foliaceum developed in the middle and upper estuary. Photosynthetic responses of phytoplankton were related to the time-scale of changes in light regime. Allocation of photosynthate to major macromolecular classes (LMWM, lipid, polysaccharide, and protein), like phytoplankton biomass and primary production, showed strong spatio-temporal variability. High carbon fixation into low molecular weight metabolites was associated with growth limitation by low light. The relative incorporation of photosynthetic carbon into proteins increased at the beginning of the phytoplankton bloom but overall, it was rather constant. However, carbon allocation into storage products such us lipid or polysaccharide increased when carbon and energy produced under optimal growth conditions exceeded what could be assimilated into protein. These patterns are explained by both spatio-temporal changes in the environmental conditions and species-specific differences. In general, daily variability appeared to be more important than diurnal periodicity in the physiological responses of phytoplankton. Results from this study show that phytoplankton photosynthesis and carbon metabolism are simultaneously affected by biotic and abiotic factors, although short-term light fluctuations may have a major influence on the physiological state of phytoplankton in the Urdaibai estuary.     

Combining a regional climate model with a phytoplankton community model to predict future changes in phytoplankton in lakes

1. Linking a regional climate model (RCM) configured for contemporary atmospheric greenhouse gas concentrations, with a phytoplankton community model (PROTECH) produced realistic simulations of 20 years of recent phytoplankton data from Bassenthwaite Lake, in the North‐West of England. 2. Meteorological drivers were derived from the RCM to represent a future climate scenario involving a 1% per annum compound increase in atmospheric CO₂ concentrations until 2100. Using these drivers, PROTECH was run for another 20 year period representing the last two decades of the 21st century. 3. Comparison of these present and future simulations revealed likely impacts on the current seasonal phytoplankton development. Under future climate conditions, the simulated spring bloom showed an increase in cyanobacteria dominance caused by greater success of Planktothrix. Also, the summer cyanobacteria bloom declined earlier because of nutrient limitation caused by the increased spring growth. Overall productivity in the lake did not change. 4. Analysis showed that these predicted changes were driven by changes in water temperature, which were in turn triggered by the higher air temperatures predicted by the RCM.     

Effects of aquatic macrophytes on water quality and phytoplankton communities in shallow lakes

We investigated aquatic macrophytes, water quality, and phytoplankton biomass and species composition in three shallow lakes with different levels of vegetation cover and nutrient concentration in Kushiro Moor, during August 2000. Trapa japonica can live in a wide range of nutrient levels. This species forms an environment with a steeper extinction of light, higher concentrations of dissolved organic carbon (DOC), lower concentrations of dissolved oxygen (DO) near the bottom, and lower concentrations of nitrate+nitrite and soluble reactive phosphorus (SRP) than other vegetation types. The pH was much higher in a Polygonum amphibium community, and the DO near the bottom did not decrease compared to a T.japonica community in the summer. The relationship between chlorophyll a and the limiting nutrient (total phosphorus (TP) when total nitrogen (TN):TPis 10 and TN/10 when TN:TP is <10) significantly differed between lakes with and without submerged vegetation. The chlorophyll a concentrations at a given nutrient level were significantly lower in water with submerged macrophytes than in water without them. Correspondence analysis showed that the difference in phytoplankton community structure across sites was largely due to the presence or absence of submerged macrophytes, and the ordination of phytoplankton species in the lakes with submerged macrophytes is best explained by environmental gradients of TN, chlorophyll, pH and SRP.     

Phytoplankton and its dynamics in two tropical lakes: a tropical and temperate zone comparison

Temporal patterns of phytoplankton biomass and community structure are described for two Kenyan lakes and subsequently compared with patterns reported in other tropical and temperate lakes. Lake Naivasha had a lower and more seasonally variable (10×) biomass, with a seasonal shift between diatoms and blue-greens, while the L. Oloidien biomass was less variable (3.7×) and dominated by blue-greens. Biomass and chlorophyll a were strongly correlated and in turn were coupled to the level of total phosphorus. A total of 143 and 94 taxa were described for L. Naivasha and L. Oloidien, respectively.The comparative analysis showed: a) a paucity of exclusively tropical species; b) that more than 30 percent of the species in two highly saline Kenyan lakes were also present in the two freshwater lakes; c) no evidence for a postulated decline of phytoplankton species abundance with latitude from the temperate zone to the tropics; d) that the low fraction of chrysophyte biomass in tropical lakes is a function of trophy rather than of latitude; e) that the fraction of chlorophyte biomass in tropical lakes is generally higher than in temperate lakes; f) that the proportion of nannoplankton in the two Kenyan freshwater lakes is not different from that in temperate lakes of the same trophy; g) that seasonal or annual biomass oscillations in the tropics are not systematically lower than in the temperate zone; h) evidence for large inter-year difference in the max.:min. biomass ratio in the only tropical lake (L. Naivasha) for which such data are available; i) that an average biomass ratio appears predictable for tropical lakes from the proportion of the sediment surface in contact with epilimnetic water. Overall, no evidence was found that the freshwater tropical phytoplankton composition or dynamics differ in any fundamental fashion from that observed in the temperate lakes during the summer.Contribution number 147 of the Limnology Research Centre, McGill University.     

Seasonal succession of phytoplankton and its diversity in two highly eutrophic lakes with different conditions of stratification

Seasonal succession of phytoplankton biomass, its diversity and its photosynthetic activity in two highly eutrophic lakes have been compared. In order to test the intermediate disturbance hypothesis, the lakes have been chosen with almost the same level of trophy but different conditions of stratification, through two ice-free periods of open water with different weather conditions.High phytoplankton diversity throughout the period of investigation was characteristic for the shallower Lake Lobardzu. The number of species here was usually more than 30 and the Shannon diversity changed from 1.2 to 4.2. Owing to the frequent external disturbances, periods characterized by autogenic succession with establishing dominance and declining diversity alternated with periods of biomass reduction and rises of diversity and photosynthetic activity. In the warmer summer of 1983, with more intense warming of bottom layers and predominance of blue-greens, phytoplankton biomass was higher and diversity lower than in the cold summer of 1982.In stratified Lake Rudusku, phytoplankton diversity and number of species were usually much lower. During the long summer stratification up to three-four dominant species of blue-greens and dinoflagellates become established and competitive exclusion leading to low diversity advanced. Some changes in biomass and diversity, were caused by zooplankton activity.     

Productivity of high-latitude lakes: climate effect inferred from altitude gradient

Climate change is predicted to be dramatic at high latitudes. Still, climate impact on high latitude lake ecosystems is poorly understood. We studied 15 subarctic lakes located in a climate gradient comprising an air temperature difference of about 6°C. We show that lake water productivity varied by one order of magnitude along the temperature gradient. This variation was mainly caused by variations in the length of the ice‐free period and, more importantly, in the supply of organic carbon and inorganic nutrients, which followed differences in terrestrial vegetation cover along the gradient. The results imply that warming will have rapid effects on the productivity of high latitude lakes, by prolongation of ice‐free periods. However, a more pronounced consequence will be a delayed stimulation of the productivity following upon changes of the lakes terrestrial surroundings and subsequent increasing input of elements that stimulate the production of lake biota.     

On the formation of net phytoplankton patches in the southern Black Sea during the spring

Species composition, community structure and spatial distribution of net phytoplankton in the southern Black Sea have been studied. In the fraction over 55 m, diatoms were found to be predominant in cell abundance and followed by dinoflagellates. Net phytoplankters were found to be more abundant in the southwestern sector, especially near the Bosporus and near Sakarya River mouth. MDS (Multi-Dimensional Scaling) analysis has projected four distinct patches along the coast, two of which appear to occupy smaller regions compared to the main western and eastern patches which have lower species numbers and diversity. The diatom species Pseudo-nitzschia seriata and Cerataulina pelagica and the dinoflagellates Noctiluca scintillans and Scrippsiella trochoidea dominate the west, while diatom species Synedra acus and Asterionella formosa together with Microspora sp. and Ankistrodesmus falcatus from chlorophyceae dominate the east.     

The impact of climate change on the physical characteristics of the larger lakes in the English Lake District

1. The larger lakes of the English Lake District have been the subject of intensive scientific study for more than 60 years. Year‐to‐year variations in the weather have recently been shown to have a major effect on their physical characteristics. The area is mild but very wet and the dynamics of the lakes are strongly influenced by the movement of weather systems across the Atlantic. 2. Here, we combine the results of long‐term measurements and the projections from a Regional Climate Model (RCM) to assess the potential impact of climate change on the surface temperature and residence times of the lakes. 3. The RCM outputs used were produced by the U.K. Hadley Centre and are based on the IPCC ‘A2’ scenario for the emission of greenhouse gases. These suggest that winters in the area will be very much milder and wetter by the 2050s and that there will be a pronounced reduction in the summer rainfall. 4. An analysis of the meteorological data acquired between 1940 and 2000 shows that there have been progressive increases in the winter air temperature and in the rainfall which are correlated with the long‐term change in the North Atlantic Oscillation. The trends reported during the summer were less pronounced and were correlated with the increased frequency of anticyclonic days and a decrease in the frequency of westerly days in the British Isles. 5. A simple model of the year‐to‐year variations in surface temperatures showed that the highest winter temperatures were recorded in the deeper lakes and the highest summer temperatures in the lakes with the shallowest thermoclines. When this model was used to predict the surface temperatures of the lakes in the 2050s, the greatest winter increase (+1.08 °C) was observed in the shallowest lake and the greatest summer increase (+2.18 °C) in the lake with the shallowest thermocline. 6. The model used to estimate the seasonal variation in the residence time of the lakes showed that the most pronounced variations were recorded in lakes with a short residence time. Average winter residence times ranged from a minimum of 10 days to a maximum of 436 days and average summer values from a minimum of 23 days to a maximum of 215 days. When this model was used to predict the residence time of the lakes in the 2050s, the greatest winter decrease (−20%) was observed in the smallest lake and the greatest summer increase (+92%) in the lake with the shortest residence time. 7. The results are discussed in relation to trends reported elsewhere in Europe and the impact of changes in the atmospheric circulation on the dynamics of the lakes. The most serious limnological effects were those projected for the summer and included a general increase in the stability of the lakes and a decrease in the flushing rate of the lakes with short residence times.     

Frost in a future climate: modelling interactive effects of warmer temperatures and rising atmospheric [CO2] on the incidence and severity of frost damage in a temperate evergreen (Eucalyptus pauciflora)

Although plants are more susceptible to frost damage under elevated atmospheric [CO₂], the importance of frost damage under future, warmer climate scenarios is unknown. Accordingly, we used a model to examine the incidence and severity of frost damage to snow gum (Eucalyptus pauciflora) in a sub‐alpine region of Australia for current and future conditions using the A2 IPCC elevated CO₂ and climate change scenario. An existing model for predicting frost effects on E. pauciflora seedlings was adapted to include effects of elevated [CO₂] on acclimation to freezing temperatures, calibrated with field data, and applied to a study region in Victoria using climate scenario data from CSIRO's Global Climate Model C‐CAM for current (1975–2004) and future (2035–2064) 30 years climate sequences. Temperatures below 0 °C were predicted to occur less frequently while the coldest temperatures (i.e. those below ?8 °C) were almost as common in the future as in the current climate. Both elevated [CO₂] and climate warming affected the timing and rates of acclimation and de‐acclimation of snow gum to freezing temperatures, potentially reducing the length of time that plants are fully frost tolerant and increasing the length of the growing season. Despite fewer days when temperatures fall below 0 °C in the future, with consequently fewer damaging frosts with lower average levels of impact, individual weather sequences resulting in widespread plant mortality may still occur. Furthermore, delayed acclimation due to either warming or rising [CO₂] combined with an early severe frost could lead to more frost damage and higher mortality than would occur in current conditions. Effects of elevated [CO₂] on frost damage were greater in autumn, while warming had more effect in spring. Thus, frost damage will continue to be a management issue for plantation and forest management in regions where frosts persist.     

Changes in the phytoplankton of Lake Trummen induced by restoration

Lake Trummen, previously an oligotrophic lake, was heavily polluted during 1930–1957. The lake was restored by suction dredging of the top sediment layer. After restoration nutrien concentrations and phytoplankton biomass were drastically reduced. Eutrophic species disappeared and more oligotrophic species returned to the lake. The investigation covers 12 years-monthly-bimonthly countings of algae together with electronmicroscopical studies of taxonomically difficult taxa. The investigation was made at the species level (300 species identified and 80 quantitatively recorded).