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).     

Light climate and phytoplankton photosynthesis in maritime Antarctic lakes

The responses of phytoplankton populations to seasonal changes in radiation flux in two Antarctic lakes with extensive winter ice-cover are described. A phytoplankton capable of photosynthesis was found throughout the year in both systems. During winter, low incident radiation combined with thick layers of snow and ice prevented in situ photosynthesis becoming detectable. The beginning of spring was marked by a reduction in snow cover which resulted in a considerable increase in surface penetrating radiation. Planktonic algae rapidly adapted to utilise these increased levels efficiently, though they still showed characteristics of strong shade adaptation.Loss of ice cover at the start of the short open water period further increased the radiation levels and a summer population developed which was much less shade adapted. Saturation and photoinhibition effects were widespread during this period as the algae proved unable to utilise high radiation levels efficiently. They were however effective at the radiation fluxes prevalent in the lower part of the rapidly circulating water columns.     

Seasonality of phytoplankton in some South Indian lakes

The landscape of South India is dotted with innumerable man-made lakes. They differ vastly in age, physiography, water flow characteristics, chemistry and trophic state, yet maintain a phytoplankton overwhelmingly dominated (43–93%) by blue-green algae; the subdominants are diatoms and/or Chlorococcales and euglenoids. The blue-greens apparently reach them from soils which are known to harbour a rich blue-green flora and several species in common with limnoplankton.South Indian lakes resemble some tropical counterparts in sustaining dense phytoplankton populations all the year round and temperate dimictic ones in showing two annual growth peaks that usually occur in summer (February–May) and the post-monsoon period (October–November), in synchrony with rise in temperature. In the chemically more oligotrophic lakes, the peaks are constituted by Raphidiopsis mediterranea Skuja, Navicula cryptocephala Kütz., Melosira granulata (Ehr.) Ralfs, and others and in hypereutrophic lakes by Microcystis aeruginosa Kütz., Synechocystis aquatilis Sauv., Oscillatoria spp., Burkillia coronata West & West and Euglena acus Ehr. The bimodal seasonality in abundance of phytoplankton reflects in chlorophyll and biomass concentrations although these are not in strict synchrony with each other. At the maxima chlorophyll a and over-dry biomass may rise to 8.5 mg l^–1 and 204 mg l^–1 respectively in highly productive waters. The highest rate of carbon assimilation recorded in such phases is 10.6 g C m^–3 d^–1.     

Spatial, seasonal and long-term variability of phytoplankton photosynthesis in lakes

A cross-system, worldwide approach has been used to ascertainthe spatial, seasonal and long-term variability of areal phytoplanktonphotosynthesis (PP) in lakes using published data sets. Also,the average fraction of annual PP occurring under ice is calculated.The lakes considered embrace a range of properties (depth, mixing,flushing rate, latitude and trophic status). The overall yearlyPP distribution is skewed to the left, suggesting the dominanceof low PP rates in the data set. When comparing lake types,no differences in average PP have been found among them. Inparticular, there are no clear areal PP differences among lakesof different trophic status on yearly, averaged basis, suggestingthat environmental limitations to PP also exist in lakes ofhigher trophic status. Volumetric-based PP can be better usedto outline PP-based trophic differences, but some degree ofoverlap is also apparent. Across all lake types (except in tropicallakes), the PP seasonal course experiences only one peak inthe year, but its timing is clearly different for each laketype. The seasonal variability of PP is lower in tropical lakes,as previously reported, but the variability of the other laketypes is roughly the same. Therefore, the effects of depth,mixing regime, flushing rate and nutrient status on PP seasonalityare difficult to ascertain since they appear to be counterbalancedby other more pervasive, local effects. Particularly, thereis no increase in temporal variability with the trophic statusof lakes, suggesting that PP seasonal control by physical variablesoverrides that of nutrients. Also, no significant relationshipbetween average PP and latitude has been found. Seasonal variabilityincreases as the yearly PP increases. On a relative basis, thereis a spatial gradient of seasonal variability of PP, which isweaker when seasonal variability of PP is considered in interyearcomparisons. Long-term (i.e. interannual) variability of PPis clearly related to increasing yearly averaged PP. Specifically,in temperate, stratifying lakes the seasonal time course ofPP is clearly different from that of phytoplankton biomass,suggesting an uncoupling of both variables as a result of differingP_max and losses throughout the year. On an average basis, environmentalvariables are poor predictors of areal daily PP, thereby implyingthat the interplay of factors is complex and changing throughoutthe year. PP under ice averages 10% of yearly PP, but its variabilityis high enough to make its measurement advisable.     

The seasonality of phytoplankton in African lakes

Although some study of the subject began in 1899, wide-ranging information from African water-bodies has only become available since 1950. Important developments included the establishment of long-term centres of research, the adoption of improved methods for quantitative algal sampling, the more intensive study of environmental conditions, the beginnings of experimental testing, and the improvement of taxonomic knowledge.At higher latitudes (> 20 °) examples of pronounced algal seasonality are long-established; they are accompanied and influenced by marked changes in radiant energy income and so water temperature, and often by effects of seasonal water input. Illustrations are given from lakes in Morocco and South Africa.More generally in Africa, including the tropical belt, annual patterns of phytoplankton seasonality are usually either dominated by hydrological features (water input-output) or by hydrographic ones (water-column structure and circulation). Examples of both types are discussed, together with instances (e.g. L. Volta) of combined hydrological and hydrographic regulation. In both the seasonal abundance of diatoms is often distinct and complementary to that of blue-green algae, with differing relationships to vertical mixing and water retention.Horizontal variability in the seasonal cycle is especially pronounced in the larger or morphometrically subdivided lakes. Some inshore-offshore differentiation is also known to affect phytoplankton quantity (e.g. L. George) and species composition (e.g. L. Victoria). Longitudinal differentiation is common in elongate basins especially when with a massive or seasonal inflow at one end (e.g. L. Turkana, L. Nubia, L. Volta); occasional terminal upwelling can also be influential (e.g. southern L. Tanganyika). Such examples grade into the longitudinally differentiated seasonality of flowing river-reservoir systems, as studied on the Blue and White Niles.The annual amplitude of population density, expressed in orders of magnitude (=log₁₀ units), is one measure of seasonal variability. It can exceed 3 orders both in systems subject to hydrological wash-out (e.g. Nile reservoirs) and in the more variable species components of lakes of long retention (e.g. L. Victoria). Low amplitudes can be characteristic of some components (e.g. green algae in L. Victoria) or of total algal biomass (e.g. L. George, L. Sibaya).Seasonal changes may be subordinated to inter-annual ones, especially in shallow and hydrologically unstable lakes (e.g. L. Nakuru).     

The influence of a dissolved inorganic nitrogen gradient on phytoplankton community dynamics in a chain of lakes

In a chain of lakes along which nutrient availability varies in a gradient, we performed factorial nutrient enrichment experiments to determine if nitrogen limitation was the principal factor controlling the differences in phytoplankton biomass, photosynthetic productivity, diversity, and species composition among two of the lakes in the chain. In the least productive lake, East Graham Lake, P and C enrichments (in the absence of N enrichment) had no effect on biomass and diversity, whereas within two weeks the N enrichments (alone or in any combination with P and/or C) increased the biomass and decreased the diversity of East Graham Lake phytoplankton to levels similar or identical to those in more productive Shoe Lake. Short-term ¹⁴C photosynthetic rates in East Graham Lake water also responded only to N in the third week. However, photosynthesis was stimulated by P in the first week, and a few species did increase in numbers with P enrichment, suggesting that some degree of P limitation remains in addition to the strong N limitation in East Graham Lake. A number of species responded individually to the enrichments in a manner similar to that of the overall community, and a strong overlapping of discriminant analysis scores for N-enriched East Graham Lake with those of Shoe Lake was consistent with our prediction that the community structure of N-enriched East Graham Lake water would shift toward that of Shoe Lake. However, many species did not respond consistently with these results, and the nutrients tested were clearly not a major factor in the differences in abundance of those species among the two lakes. The results support the argument that overall biomass production and diversity of the phytoplankton community in a lake can be a relatively simple function of a single most-limiting nutrient. However, many of the species responses also confirm that, while nutrient availability is an important factor in the control of the species composition of the community, other factors are likely to prevent reliable predictions of all species effects on the basis of nutrient availability alone.     

The interaction of components controlling net phytoplankton photosynthesis in a well-mixed lake (Lough Neagh, Northern Ireland)

The homogeneous distribution of the phytoplankton in a shallow (mean depth 8·6 m) unstratified lake, L. Neagh, Northern Ireland, facilitated the study of the interaction of components controlling gross photosynthesis per unit area. These included the photosynthetic capacity, the phytoplankton content of the euphotic zone, and a logarithmic function describing the effective radiation input. These factors were analysed for two sites, the open lake and Kinnego Bay, which respectively had standing crops of up to 90 and 300 mg chlorophyll a m^?3 and maximum daily rates of gross integral photosynthesis of 11·7 and 15·6 g O₂ m^?2 day^?1. Values are reduced by the high contribution to light attenuation by non-algal sources, which increases at low standing crops particularly in winter, when values of integral photosynthesis decrease to 0·5 g O₂ m^?2 day^?1. This relative change is the result of self-shading behaviour of the phytoplankton altering the crop content of the euphotic zone at different population densities. Changes in the irradiance function, incorporating day length, are largely responsible for the changes in daily rates of integral gross photosynthesis; as daily irradiance is also a determinant of water temperature, it exerts further influence through the photosynthetic capacity which was strongly correlated with temperature. Much of the gain in gross photosynthesis resulting from higher photosynthetic capacity may not be reflected in a higher net column photosynthesis, because of the greater proportional rise in respiration with temperature. The balance in the water column between respiration losses and photosynthetic input may frequently alter since the ratio of illuminated to dark zones is between 1/4 to 1/5 in the open lake, and small shifts in any of the controlling features may result in conditions unfavourable for growth. This is analysed especially for the increase of diatoms in spring, when small modifications of the underwater light field can delay growth.     

The role of phytoplankton photosynthesis in global biogeochemical cycles

Phytoplankton biomass in the world's oceans amounts to only 1–2% of the total global plant carbon, yet these organisms fix between 30 and 50 billion metric tons of carbon annually, which is about 40% of the total. On geological time scales there is profound evidence of the importance of phytoplankton photosynthesis in biogeochemical cycles. It is generally assumed that present phytoplankton productivity is in a quasi steady-state (on the time scale of decades). However, in a global context, the stability of oceanic photosynthetic processes is dependent on the physical circulation of the upper ocean and is therefore strongly influenced by the atmosphere. The net flux of atmospheric radiation is critical to determining the depth of the upper mixed layer and the vertical fluxes of nutrients. These latter two parameters are keys to determining the intensity, and spatial and temporal distributions of phytoplankton blooms. Atmospheric radiation budgets are not in steady-state. Driven largely by anthropogenic activities in the 20th century, increased levels of IR- absorbing gases such as CO₂, CH₄ and CFC's and NO_x will potentially increase atmospheric temperatures on a global scale. The atmospheric radiation budget can affect phytoplankton photosynthesis directly and indirectly. Increased temperature differences between the continents and oceans have been implicated in higher wind stresses at the ocean margins. Increased wind speeds can lead to higher nutrient fluxes. Throughout most of the central oceans, nitrate concentrations are sub-micromolar and there is strong evidence that the quantum efficiency of Photosystem II is impaired by nutrient stress. Higher nutrient fluxes would lead to both an increase in phytoplankton biomass and higher biomass-specific rates of carbon fixation. However, in the center of the ocean gyres, increased radiative heating could reduce the vertical flux of nutrients to the euphotic zone, and hence lead to a reduction in phytoplankton carbon fixation. Increased desertification in terrestrial ecosystems can lead to increased aeolean loadings of essential micronutrients, such as iron. An increased flux of aeolean micronutrients could fertilize nutrient-replete areas of the open ocean with limiting trace elements, thereby stimulating photosynthetic rates. The factors which limit phytoplankton biomass and photosynthesis are discussed and examined with regard to potential changes in the Earth climate system which can lead the oceans away from steady-state. While it is difficult to confidently deduce changes in either phytoplankton biomass or photosynthetic rates on decadal time scales, time-series analysis of ocean transparency data suggest long-term trends have occurred in the North Pacific Ocean in the 20th century. However, calculations of net carbon uptake by the oceans resulting from phytoplankton photosynthesis suggest that without a supply of nutrients external to the ocean, carbon fixation in the open ocean is not presently a significant sink for excess atmospheric CO₂.The submitted paper has been authored under Contract No. DE-AC02-76H00016 with the US Department of Energy. Accordingly, the US Government retains a non-exclusive, royalty-free license to publish or reproduce the published form of this contribution, or allow others to do so, for US Government purposes.     

The influence of different disturbance frequencies on the species richness,diversity and equitability of phytoplankton in shallow lakes

The relationships between the species richness, diversity and equitability of phytoplankton is discussed in the context of Connell's (1978, Science 199: 1304–1310) Intermediate Disturbance Hypothesis (IDH). The records of 759 vertical phytoplankton samples, which were obtained from four shallow central European lakes (Balaton, Neusiedlersee, and two small artificial ponds) at daily to weekly intervals were analysed.

1. The Shannon-Weaver function was used to measure diversity of the recorded species compositionof the phytoplankton. It is shown on fictitious data that compositional diversity is sensitive to the numberof coequilibrating species provided that the suspected interrelationship between diversity and ‘complexity’is amenable to the application of this method.
2. The disturbance scale that was developed on the basis of the field records fits well to Reynolds'(1988, Verh. int. Ver. Limnol. 23: 683–691) derivation: < 3 days qualifies as high frequency, approximatel3–8 days as intermediate frequency and > 8–9 days as low frequency of disturbance for phytoplankton.
3. Arithmetical means of the compositional diversity of phytoplankton under different frequencies ofdisturbance support the hypothesis that maximal diversity appears at intermediate frequencies.
4. There are different reasons for decrease in diversity at higher and lower frequencies. Inequitabilitydiminishes diversity at low disturbance; while species number decreases at high frequencies.
5. The case of Neusiedlersee calls attention to the fact that it is difficult, if at all possible, to differentiatebetween the indices under continuous stress and high frequency of disturbance in lakes in temperateregions. Similar species number-equitability pattern are induced by both and it is also presumablethat high frequency disturbance can itself effect a serious stress.
6. The striking effects that regular major periodic events (e.g. significant changes in the grazing pressureat the onset of the clear-water phase, autumnal cooling) in the plankton have on its species diversity areevident. Thus, the relative importance of intermediate frequency disturbances has its own seasonality:it is increasingly important in periods (partly in the spring, but mostly in the summer-autumn equilibriumphases), in which competition among phytoplankton species is increasing. This observation suggestsa way by which the stochasticity-based IDH can be incorporated into rather more deterministicexplanations (e.g. PEG-model; Sommer et al., 1986. Archiv für Hydrobiologie 106: 433–471) of planktonsuccession.
7. The most controversial issue and, therefore, the main difficulty, with IDH is that it not onlymaintains species richness in an ecosystem but it also supposes its presence. The lack of either earlyor late successional species in a given community can inactivate the mechanism. From the point of viewof the diversity-species richness relationship, the persistence of disturbance at given frequencies is ofgreater importance than the temporal alterations themselves in the evolutionary ecology of the phytoplankton.
8. For characteristically unperturbed phytoplankton communities (no case was studied here), equilibriumconcepts (niche diversification, etc.) should be more strongly applicable to their diversity andspecies richness.

     

Effect of temperature on rate of photosynthesis in Antarctic phytoplankton

Summary The rate of photosynthesis of marine antarctic phytoplankton (western Scotia Sea and Bransfield Strait) was determined as a function of temperature, from ambient (-0.8°C to 1.0°C) to 28°C. Photosynthetic rates, based on radiocarbon incorporation during half-day incubations, were increased by as much as 2x with temperatures up to 7°C; at higher temperatures the rates decreased rapidly, so that at 28°C the rates were only 3% of that at ambient temperatures. In antarctic surface waters during the austral summer the rate of photosynthesis by phytoplankton thus is limited by thermodynamic effects on metabolic reactions, in spite of high nutrient concentrations and saturating light levels. The observed rates were in agreement with thermodynamic models of the dependence of phytoplankton growth rate on temperature.     

Phytoplankton photosynthesis parameters in central Canadian lakes

Values for two biological parameters — (i) P^B_m, the lightsaturated rate of photosynthesis per unit of chlorophyll and(ii) , the slope of the light limited part of the photosynthesisversus light curve per unit of chlorophyll — must be knownin order to be able to estimate rates of phytoplankton primaryproduction from chlorophyll data. These parameters were measuredfor periods of up to 9 years in central Canadian lakes locatedin temperate, subarctic and arctic climatic zones. Regardlessof their geographic locations, lakes where integral photosynthesiswas nutrient limited had lower values of these parameters thandid lakes where integral photosynthesis was light limited. Temperatureset an upper limit to the variability of P^B_m but was not a goodpredictor of its actual value. Year-to-year variability of photosynthesisparameters in the most intensively studied group of lakes waslarge: annual means varied by a factor of three over a 9-yearperiod. Until the sources and extent of this variability areknown, accurate production estimates can be obtained from chlorophylldata only if P^B_m and are measured in each water body and inevery year. Implications for estimating primary production fromremotely-sensed chlorophyll data are discussed.     

Inorganic carbon limitation of photosynthesis in lake phytoplankton

1. Inorganic carbon availability influences species composition of phytoplankton in acidic and highly alkaline lakes, whereas the overall influence on community photosynthesis and growth is subject to debate.
2. The influence of total dissolved inorganic carbon (DIC) and free CO₂ on community photosynthesis was studied in six Danish lakes during the summer of 1995. The lakes were selected to ensure a wide range of chlorophyll a concentrations (1–120 μg l^–1), pH (5.6–9.6) and DIC concentration (0.02–2.5 m m ). Photosynthesis experiments were performed using the ¹⁴C technique in CO₂-manipulated water samples, either by changing the pH or by adding/removing CO₂.
3. Lake waters were naturally CO₂ supersaturated during most of the experimental period and inorganic carbon limitation of photosynthetic rates did not occur under ambient conditions. However, photosynthesis by phytoplankton in lakes with low and intermediate DIC concentrations was seriously restricted when CO₂ concentrations declined. Similarly, photosynthesis was limited by low CO₂ concentrations during phytoplankton blooms in the hardwater alkaline lakes.     

The influence of chilling on photosynthesis and activities of some enzymes of sucrose metabolism in Lycopersicon esculentum Mill

The effects of chilling stress on leaf photosynthesis and sucrose metabolism were investigated in tomato plants (Lycopersicon esculentum Mill. cultivar Marmande). Twenty-one-day-old seedlings were grown in a growth chamber at 25/23 °C (day/night) (control) and at 10/8 °C (day/night) (chilled) for 7 days. The most evident effect of chilling was the marked reduction of plant growth and of CO₂ assimilation as measured after 7 days, the latter being associated with a decrease in stomatal closure and an increase in Ci. The inhibition in photosynthetic rate was also related to an impairment of photochemistry of photosystem II (PSII), as seen from the slight, but significant change in the ratio of F_v/F_m. The capacity of chilled leaves to maintain higher q_P values with respect to the controls suggests that some protection mechanism prevented excess reduction of PSII acceptors. The results of the determination of starch and soluble sugar content could show that chilling impaired sucrose translocation. The activity of leaf invertase increased significantly in chilled plants, while that of other sucrose-metabolizing enzymes was not affected by growing temperature. Furthermore, the increase in invertase (neutral and acid) activity, which is typical of senescent tissue characterized by reduced growth, seems to confirm that tomato is a plant which is not a plant genetically adapted to low temperatures.     

The impact of climate on the geographical distribution of phytoplankton species in boreal lakes

Here, we use a novel space-by-time approach to study large-scale changes in phytoplankton species distribution in Swedish boreal lakes in response to climate variability. Using phytoplankton samples from 27 lakes, evenly distributed across Sweden, all relatively unimpacted by anthropogenic disturbance and sampled annually between 1996 and 2010, we found significant shifts in the geographical distribution of 18 species. We also found significant changes in the prevalence of 45 species (33 became more common and 12 less common) over the study period. Using species distribution models and phytoplankton samples from 60 lakes sampled at least twice between 1992 and 2010, we evaluated the importance of climate variability and other environmental variables on species distribution. We found that temperature (e.g., extreme events and the duration of the growing season) was the most important predictor for species detections. Many cyanobacteria, chlorophytes, and, to a lesser extent, diatoms and zygnematophytes, showed congruent and positive responses to temperature. In contrast, precipitation explained little variation and was important only for a few taxa (e.g., Staurodesmus spp., Trachelomonas volvocina). At the community level, our results suggest a change in community composition at temperatures over 20 °C and growing seasons longer than 40 days. We conclude that climate is an important driver of the distributional patterns of individual phytoplankton species and may drive changes in community composition in minimally disturbed boreal lakes.     

The influence of landscape position on lakes in northern Wisconsin

1. Using data from the North Temperate Lakes Long-Term Ecological Research site in northern Wisconsin, we present a series of examples illustrating how landscape setting can influence the static and dynamic aspects of many physical, chemical and biological properties of lakes. 2. One important landscape attribute is the hydrologic position of a lake within the regional flow regime. Lake position determines the relative importance of groundwater and precipitation input to a lake, with lakes high in the landscape receiving a greater proportion of their input waters from precipitation than lakes lower in the landscape. Landscape position is strongly correlated with the concentration of base cations such as calcium and magnesium. 3. Landscape position also influences how lakes respond to drought conditions. Lakes high in the landscape responded to a 4-year drought with decreases in calcium mass, whereas lakes low in the landscape increased in mass of calcium. During extended dry conditions, these differential responses of lakes suggest that lakes already low in calcium (i.e. in a high position in the flow system) will have further reductions in calcium concentrations. These reductions could decrease the number of lakes offering suitable habitat for organisms such as crayfish and snails whose distributions are limited by calcium. 4. Landscape position also affects silica concentrations in lakes, with lakes low in the landscape having silica concentrations up to three orders of magnitude greater than lakes high in the landscape. Differences in silica concentration affect robustness of freshwater sponge spicules which can potentially alter some aspects of the dynamics of littoral zone food webs. 5. Landscape position can influence the vertical distribution of primary production. Concentrations of dissolved organic carbon are affected by landscape setting and can influence vertical light penetration, thus affecting the depth at which primary production can occur. 6. Lake area and fish species richness are correlated with landscape position: larger, species-rich lakes are low in the landscape, whereas smaller lakes with fewer species tend to be high in the landscape. 7. By taking a landscape-scale view, in addition to the more usual lake-specific view, it is possible to reach a more robust understanding of lake dynamics and avoid some of the problems associated with extrapolating from single lake results.     

Spring phytoplankton of 54 small lakes in southern Finland

The abundance and species composition of phytoplankton communities were studied rapidly following the spring ice-melt in 54 small Finnish lakes that form a unique mosaic of water bodies. Phytoplankton biomass and cell density varied among the study lakes with a factor 100 between the lowest and highest values. Highest biomass and densities of phytoplankton characterized small ( < 0.05 km²) lakes with moderate or high water colour (> 80 mg Pt l^–1). In contrast, biomass was low in clear-water lakes and lakes where water throughflow was strong. Typically one species dominated most phytoplankton communities, and usually comprised up to about 45% of the total phytoplankton biomass. Two-thirds of the 103 taxa observed were Chrysophyceans and Chlorophyceans. The most common taxa wereChlamydomonas spp. (Chlorophyceae) andCryptomonas ovata (Cryptophyceae).