Interactions between nutrient availability and climatic fluctuations as determinants of the long-term phytoplankton community changes in Lake Garda,Northern Italy

Major focus in interpreting phytoplankton changes in specific typologies of waterbodies or in single lakes is directed towards nutrients and climatic dynamics. During the last 35 years, Lake Garda (Northern Italy; A = 368 km², z _max = 350 m, V = 49 km³) underwent a significant increase of phosphorus in the water column, from ca. 10 μg P l⁻¹ to 18–22 μg P l⁻¹. At the multi-decadal scale, the increase of the trophic status had a positive impact on the growth of Cyanobacteria (mainly Oscillatoriales) and, partly, diatoms, as demonstrated by the long-term ecological research carried out since the beginning of the 1990s in the deepest zone of the lake. Conversely, the increase of Peridiniales (mostly Ceratium hirundinella) in the recent years appeared also associated with the interannual variations of lake temperature. At the seasonal and annual scale, the development of the large diatoms and Oscillatoriales during the periods of their maximum growth (early spring, and summer and autumn, respectively) was strongly controlled by the extent of spring vertical water mixing and nutrient fertilization of surface waters, which, in turn, were negatively dependent on the air and water temperatures in winter and early spring. Therefore, contrary to the positive impact of milder winters on phytoplankton growth in many lakes of high latitudes, warmer winter temperatures in deep oligomictic lakes of lower latitudes can determine periodic shifts towards more oligotrophic conditions and a minor development of diatoms and specific harmful cyanobacterial groups (Oscillatoriales). The complex relationships between the explanatory and response variables were tested by applying Path Analysis (Structural Equation Modeling). This multiequational technique has great potential for studying causal relationships in temporally ordered variables. The results highlight the necessity to study the consequences of climatic fluctuations on the phytoplankton communities at different temporal scales and complexity, also including the indirect effects of climatic dynamics mediated by the morphometric, morphological and hydrological characteristics of lakes, and the possible synergic or opposite effects with other forcing variables, including nutrients.     

Factors affecting the seasonality and distribution of cyanobacteria and chlorophytes: a case study from the large lakes south of the Alps,with special reference to Lake Garda

Chlorophytes and cyanobacteria are among the most typical algal groups, contributing to the aesthetic appearance and quality of the epilimnetic waters of the deep (251–410 m) and large (6.5–49×10⁹ m³) lakes located on the southern edge of the Alps (from oligo-mesotrophy to meso-eutrophy: Maggiore, Garda, Como, Iseo and Lugano). The results obtained from monthly surveys carried out in the largest of these lakes (Garda) have been reported in detail. The thermal stability of the water column and silica depletion were the main factors responsible for the decline of the great spring diatoms. The successive growth of Mougeotia sp. was favoured by its lower sinking velocity and resistance to the increasing grazing pressure. During summer, the maximum stability of the water column, with high vertical nutrient concentration gradients, determined a major algal differentiation with a typical increase, among others, of Chlorococcales at the surface and metalimnetic stratification of various Oscillatoriales. The development of oligotrophic blooms, caused by a rapid thickening at the surface of Anabaena in the eastern, sheltered basin, was also discussed in light of the trophic characterisation of Lake Garda. From autumn to spring, the decreasing light, the increasing mixing depth and nutrient availability favoured a progressive dominance of vertical homogeneous populations of Planktothrix (autumn) and colonial diatoms. The same functional groups of chlorophytes and cyanobacteria have been recognised in the studies published so far on the phytoplankton of deep southern subalpine lakes. In this respect, their morphometric and physical properties appear to constitute a sort of standardising factor, reducing the range of possible dominants. With increasing TP concentrations and biomass, filaments of Planktothrix and Planktolyngbya, along with the Chlorococcales, became important. The dominance of Mougeotia (one of the most characteristic features of these large lakes) appears restricted to a medium trophic range, whereas the distribution of the Chroococcales and other filaments ascribed to Pseudanabaena and/or Limnothrix is more irregular. The conspicuous presence of Aphanizomenon in Lake Lugano is typical and characteristic of a more productive lake. Among the Nostocales, a clear interpretation of the Anabaena blooms in lakes Garda and Iseo is complicated by the peculiar behaviour of filaments concentrating at the surface, which is apparently restricted, within the medium trophic range, to stable water columns.     

Long-term trends and fine year-to-year tuning of phytoplankton in large lakes are ruled by eutrophication and atmospheric modes of variability

This study demonstrated how the impact of eutrophication in a deep lake at the southern border of the Alps (Lake Garda) was regulated by specific modes of atmospheric circulation relevant for the Mediterranean area. At the decadal scale, nutrients and phytoplankton increased concurrently since the 1970s. At the annual scale, year-to-year fluctuations in nutrients and phytoplankton were controlled through a chain of causal factors centred on deeply penetrative mixing events determining an upward transport of phosphorus from the hypolimnion to the trophogenic layers. The extent of mixing was in turn controlled by lake and air winter temperature, which were ultimately regulated by the winter fluctuations of the East Atlantic pattern (EA). In its negative state, the EA shows an intense high pressure over the West Atlantic, causing a north-easterly air flow bringing cold air from continental Europe to Mediterranean, thus favouring greater lake mixing and nutrient fertilisation. Cyanobacteria (mostly Planktothrix rubescens) were the organisms which greatly benefitted from the long-term increase in phosphorus concentrations and the year-to-year fluctuations in surface phosphorus availability controlled by the EA. Given the same availability of phosphorus in the water column, positive winter EA phases weakened the eutrophication effects and phytoplankton development.     

Weekly dynamics of phytoplankton functional groups under high water level fluctuations in a subtropical reservoir-bay

In reservoirs, water level fluctuations strongly influence phytoplankton development. However, studies on the response of phytoplankton in the reservoir-bay to water level fluctuations are very scarce, especially in the highly dynamic reservoir system, for instance, the Three Gorges Reservoir (TGR) on the Yangtze River in China. Therefore, we carried out weekly monitoring in a typical tributary bay—Xiangxi Bay of the TGR from March 2008 to March 2009, to analyze the dynamics of phytoplankton functional groups, as well as their response to the water level fluctuations and other environmental conditions. The phytoplankton functional groups G (short, nutrient-rich water columns with high light and without nutrient deficiency), M (dielly mixed layers of small eutrophic, low latitude with high insolation and without flushing and low total light) and Lo (summer epilimnia in mesotrophic lakes with segregated nutrients and without prolonged or deep mixing) were the most important in biomass, mainly represented by Pandorina morum and Eudorina elegans, Microcystis aeruginosa, Peridiniopsis niei and Ceratium hirundinella, respectively. The dominant functional groups had close relationships with the water level fluctuations, light and nutrient, etc. Principal components analysis and redundancy analysis indicated that phytoplankton functional groups in Xiangxi Bay were restricted by the mixing regime and other abiotic variables under the influences of the mixing regime. In Xiangxi Bay, the water level fluctuation showed significant correlations with many physicochemical variables, including the mixing depth (r = 0.97, p < 0.001) and the relative water column stability (r = −0.80, p < 0.001). The study implied that water level fluctuations had complex influence on environmental changes and selecting for phytoplankton functional groups in a highly dynamic reservoir-bay. The important characteristics of the dominant phytoplankton functional groups in Xiangxi Bay were also discussed.     

Long-term response of a shallow, moderately flushed lake to reduced external phosphorus and nitrogen loading

1. The responses of nutrient concentrations, plankton, macrophytes and macrozoobenthos to a reduction in external nutrient loading and to contemporary climatic change were studied in the shallow, moderately flushed Lake Müggelsee (Berlin, Germany). Weekly to biweekly data from 1979 to 2003 were compared with less frequently collected historical data. 2. A reduction of more than 50% in both total phosphorus (TP) and total nitrogen (TN) loading from the hypertrophic (1979–90) to the eutrophic period (1997–2003) was followed by an immediate decline in TN concentrations in the lake. TP concentrations only declined during winter and spring. During summer, phosphorus (P) release from the sediments was favoured by a drastic reduction in nitrate import. Therefore, Müggelsee acted as a net P source for 6 years after the external load reduction despite a mean water retention time of only 0.1–0.16 years. 3. Because of the likely limitation by P in spring and nitrogen (N) in summer, phytoplankton biovolume declined immediately after nutrient loading was reduced. The formerly dominant cyanobacteria (Oscillatoriales) Limnothrix redekei and Planktothrix agardhii disappeared, but the mean biovolume of the N₂‐fixing species Aphanizomenon flos‐aquae remained constant. 4. The abundance of Daphnia spp. in summer decreased by half, while that of cyclopoid copepod species increased. Abundances of benthic macroinvertebrates (mainly chironomids) decreased by about 80%. A resource control of both phytoplankton and zooplankton is indicated by significant positive correlations between nutrient concentrations and phytoplankton biovolume and between phytoplankton and zooplankton biomass. 5. Water transparency in spring increased after nutrient reduction and resulted in re‐colonisation of the lake by Potamogeton pectinatus. However, this process was severely hampered by periphyton shading and grazing by waterfowl and fish. 6. Water temperatures in Müggelsee have increased in winter, early spring and summer since 1979. The earlier development of the phytoplankton spring bloom was associated with shorter periods with ice cover, while direct temperature effects were responsible for the earlier development of the daphnid maximum in spring.     

The recovery of a very shallow eutrophic lake, 20 years after the control of effluent derived phosphorus

1. Monitoring at fortnightly to monthly intervals of a very shallow, lowland lake over 24 years has enabled the time course of recovery from nutrient enrichment to be investigated after high external P loading of the lake (>10 g P m^?2 year^?1) was reduced between 1977 and 1980. 2. The lake showed a relatively rapid response during the spring and early summer, with a reduction in phytoplankton biomass occurring after 5 years when soluble reactive phosphorus concentration was <10 μg L^?1. 3. However, during the later summer the response was delayed for 15 years because of sustained remobilisation of phosphorus from the sediment. The greater water clarity in spring and a gradual shift from planktonic to benthic algal growth may be related to the reduction in internal loading after 15 years. 4. Changes in the phytoplankton community composition were also observed. Centric diatoms became less dominant in the spring, and the summer cyanobacteria populations originally dominated by non‐heterocystous species (Limnothrix/Planktothrix spp.) almost disappeared. Heterocystous species (Anabaena spp. and Aphanizomenon flos‐aquae) were slower to decline, but after 20 years the phytoplankton community was no longer dominated by cyanobacteria. 5. There were no substantial changes in food web structure following re‐oligotrophication. Total zooplankton biomass decreased but body size of Daphnia hyalina, the largest zooplankton species in the lake, remained unchanged, suggesting that the fish population remained dominated by planktivorous species. 6. Macrophyte growth was still largely absent after 20 years, although during the spring water clarity may have become sufficient for macrophytes to re‐establish.     

Water temperature and stratification depth independently shift cardinal events during plankton spring succession

In deep temperate lakes, the beginning of the growing season is triggered by thermal stratification, which alleviates light limitation of planktonic producers in the surface layer and prevents heat loss to deeper strata. The sequence of subsequent phenological events (phytoplankton spring bloom, grazer peak, clearwater phase) results in part from coupled phytoplankton–grazer interactions. Disentangling the separate, direct effects of correlated climatic drivers (stratification‐dependent underwater light climate vs. water temperature) from their indirect effects mediated through trophic feedbacks is impossible using observational field data, which challenges our understanding of global warming effects on seasonal plankton dynamics. We therefore manipulated water temperature and stratification depth independently in experimental field mesocosms containing ambient microplankton and inocula of the resident grazer Daphnia hyalina. Higher light availability in shallower surface layers accelerated primary production, warming accelerated consumption and growth of Daphnia, and both factors speeded up successional dynamics driven by trophic feedbacks. Specifically, phytoplankton peaked and decreased earlier and Daphnia populations increased and peaked earlier at both shallower stratification and higher temperature. The timing of ciliate dynamics was unrelated to both factors. Volumetric peak densities of phytoplankton, ciliates and Daphnia in the surface layer were also unaffected by temperature but declined with stratification depth in parallel with light availability. The latter relationship vanished, however, when population sizes were integrated over the entire water column. Overall our results suggest that, integrated over the entire water column of a deep lake, surface warming and shallower stratification independently speed up spring successional events, whereas the magnitudes of phytoplankton and zooplankton spring peaks are less sensitive to these factors. Therefore, accelerated dynamics under warming need not lead to a trophic mismatch (given similar grazer inocula at the time of stratification). We emphasize that entire water column dynamics must be studied to estimate global warming effects on lake ecosystems.     

Mechanisms preventing a decrease in phytoplankton biomass after phosphorus reductions in a German drinking water reservoir—results from more than 50 years of observation

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Phytoplankton dynamics and structure: a comparative analysis in natural and man-made water bodies of different trophic state

Previous investigations on Sicilian man made lakes suggested that physical factors, along with the specific morphology and hydrology of the water body, are important in selecting phytoplankton species. In particular, the variations of the z _mix/z _eu ratio due to the operational procedure to which reservoirs are generally subject were recognised as a trigger allowing the assemblage shift. To investigate if these variations may be considered analogous to those occurring in natural lakes as trophic state and phytoplankton biomass increase, causing a transparency decrease and a contraction of the euphotic depth, phytoplankton were collected in two natural water bodies, one mesotrophic (Lake Biviere di Cesarò) the other eutrophic (Lake Soprano), and compared with those collected in two reservoirs with analogous trophic characteristics (Lake Rosamarina, mesotrophic and Lake Arancio, eutrophic). Particular attention was paid to the dynamics of two key groups: Cyanophytes and chlorophytes. In all four water bodies, transparency mainly depended on chlorophyll level. Annual average value of phytoplankton biomass in the mesotrophic environments was below 2.0 mg l^–1, whereas in the eutrophic systems it was well above 10 mg l^–1. All water bodies showed the presence of cyanophytes (e.g. Anabaena spp., Anabaenopsis spp., Microcystis spp., Planktothrix spp.) and chlorophytes (e.g. Chlamydomonas spp., Botryococcus spp., Oocystis spp., Scenedesmus spp., Pediastrum spp.), but their relative proportions and body size dimensions were different. In particular, small colonial chlorophytes and large-colony forming cyanophytes were most common in the most eutrophic water bodies, whereas larger colonies of green algae in those with a lower trophic state. The results showed that, under the same climatic conditions, autogenic (increase of biomass, decrease in light penetration and euphotic depth) and allogenic (use of the stored waters, anticipated breaking of the thermocline, increase of the mixing depth) processes may shift the structure of phytoplankton assemblage in the same direction even though the quantity of biomass remains linked to nutrient availability.     

TN : TP ratio and planktivorous fish do not affect nutrient-chlorophyll relationships in shallow lakes

1. In previous work, phytoplankton regulation in freshwater lakes has been associated with many factors. Among these, the ratio of total nitrogen to total phosphorus (TN : TP) has been widely proposed as an index to identify whether phytoplankton are N‐ or P‐limited. From another point of view, it has been suggested that planktivorous fish can be used to control phytoplankton. 2. Large‐scale investigations of phytoplankton biomass [measured as chlorophyll a, (chl‐a)] were carried out in 45 mid‐lower Yangtze shallow lakes to test hypotheses concerning nutrient limitation (assessed with TN : TP ratios) and phytoplankton control by planktivorous fish. 3. Regression analyses indicated that TP was the primary regulating factor and TN the second regulating factor for both annual and summer phytoplankton chl‐a. In separate nutrient–chl‐a regression analyses for lakes of different TN : TP ratios, TP was also superior to TN in predicting chl‐a at all particular TN : TP ranges and over the entire TN : TP spectrum. Further analyses found that chl‐a : TP was not influenced by TN : TP, while chl‐a : TN was positively and highly correlated to TP : TN. 4. Based on these results, and others in the literature, we argue that the TN : TP ratio is inappropriate as an index to identify limiting nutrients. It is almost impossible to specify a ‘cut‐off’ TN : TP ratio to identify a limiting nutrient for a multi‐species community because optimal N : P ratios vary greatly among phytoplankton species. 5. Lakes with yields of planktivorous fish (silver and bighead carp, the species native to China) >100 kg ha^?1 had significantly higher chl‐a and lower Secchi depth than those with yields <100 kg ha^?1. TP–chl‐a and TP–Secchi depth relationships are not significantly different between lakes with yields >100 kg ha^?1 or <100 kg ha^?1. These results indicate that the fish failed to decrease chl‐a yield or enhance Z_SD. Therefore, silver carp and bighead carp are not recommended as a biotic agent for phytoplankton control in lake management if the goal is to control the entire phytoplankton and to enhance water quality.     

Do phytoplankton communities correctly track trophic changes? An assessment using directly measured and palaeolimnological data

1. Measurements of total phosphorus (TP) concentrations since 1975 and a 50‐year time series of phytoplankton biovolume and species composition from Lake Mondsee (Austria) were combined with palaeolimnological information on diatom composition and reconstructed TP‐levels to describe the response of phytoplankton communities to changing nutrient conditions. 2. Four phases were identified in the long‐term record. Phase I was the pre‐eutrophication period characterised by TP‐levels of about 6 μg L^?1 and diatom dominance. Phase II began in 1966 with an increase in TP concentration followed by the invasion of Planktothrix rubescens in 1968, characterising mesotrophic conditions. Phase III, from 1976 to 1979, had the highest annual mean TP concentrations (up to 36 μg L^?1) and phytoplankton biovolumes (3.57 mm³ L^?1), although reductions in external nutrient loading started in 1974. Phases II and III saw an expansion of species characteristic of higher nutrient levels as reflected in the diatom stratigraphy. Oligotrophication (phase IV) began in 1980 when annual average TP concentration, Secchi depth and algal biovolume began to decline, accompanied by increasing concentrations of soluble reactive silica. 3. The period from 1981 to 1986 was characterised by asynchronous trends. Annual mean and maximum total phytoplankton biovolume initially continued to increase after TP concentration began to decline. Reductions in phytoplankton biovolume were delayed by about 5 years. Several phytoplankton species differed in the timing of their responses to changing nutrient conditions. For example, while P. rubescens declined concomitantly with the decline in TP concentration, other species indicative of higher phosphorus concentrations, such as Tabellaria flocculosa var. asterionelloides, tended to increase further. 4. These data therefore do not support the hypotheses that a reduction in TP concentration is accompanied by (i) an immediate decline in total phytoplankton biovolume and (ii) persistence of the species composition characterising the phytoplankton community before nutrient reduction.     

Effects of fertiliser and crayfish on plankton and nutrient dynamics in hardwater ponds

Although phosphorus fertilisation can improve productivity in most freshwater ponds, phosphate may become limiting in extremely hard water due to rapid precipitation with calcium. Hence we studied the characteristics of plankton and nutrient dynamics in water containing >400 mg CaCO₃ l^–1in pond and microcosm systems. The field experiment was conducted in eight earthen ponds involving two nutrient ratios (N:P = 1:1 and 20:1) with or without crayfish. Fertilisation significantly increased concentrations of NO₂–N and NO₃–N, but soluble reactive phosphorus was depleted to the level prior to fertilisation within 24 h. The laboratory test showed that after 6 h of fertilisation, 45% phosphorus was precipitated by calcium, 30% phosphorus was assimilated by phytoplankton and only 25% phosphorus remained in water column. The phytoplankton abundance in hardwater ponds was regulated by the abundance of zooplankton population rather than by either crayfish or fertilisation. The presence of crayfish only increased the concentration of total phosphorus. This study suggests that when phytoplankton production is required in crayfish ponds the maintenance of phytoplankton abundance will depend on the effective control of zooplankton rather than fertilisation. Due to the rapid precipitation of phosphorus by calcium in hard water ponds, more frequent phosphorus fertilisation is needed to enhance primary productivity.     

Lake responses to reduced nutrient loading – an analysis of contemporary long-term data from 35 case studies

1. This synthesis examines 35 long‐term (5–35 years, mean: 16 years) lake re‐oligotrophication studies. It covers lakes ranging from shallow (mean depth <5 m and/or polymictic) to deep (mean depth up to 177 m), oligotrophic to hypertrophic (summer mean total phosphorus concentration from 7.5 to 3500 μg L^?1 before loading reduction), subtropical to temperate (latitude: 28–65°), and lowland to upland (altitude: 0–481 m). Shallow north‐temperate lakes were most abundant. 2. Reduction of external total phosphorus (TP) loading resulted in lower in‐lake TP concentration, lower chlorophyll a (chl a) concentration and higher Secchi depth in most lakes. Internal loading delayed the recovery, but in most lakes a new equilibrium for TP was reached after 10–15 years, which was only marginally influenced by the hydraulic retention time of the lakes. With decreasing TP concentration, the concentration of soluble reactive phosphorus (SRP) also declined substantially. 3. Decreases (if any) in total nitrogen (TN) loading were lower than for TP in most lakes. As a result, the TN : TP ratio in lake water increased in 80% of the lakes. In lakes where the TN loading was reduced, the annual mean in‐lake TN concentration responded rapidly. Concentrations largely followed predictions derived from an empirical model developed earlier for Danish lakes, which includes external TN loading, hydraulic retention time and mean depth as explanatory variables. 4. Phytoplankton clearly responded to reduced nutrient loading, mainly reflecting declining TP concentrations. Declines in phytoplankton biomass were accompanied by shifts in community structure. In deep lakes, chrysophytes and dinophytes assumed greater importance at the expense of cyanobacteria. Diatoms, cryptophytes and chrysophytes became more dominant in shallow lakes, while no significant change was seen for cyanobacteria. 5. The observed declines in phytoplankton biomass and chl a may have been further augmented by enhanced zooplankton grazing, as indicated by increases in the zooplankton : phytoplankton biomass ratio and declines in the chl a : TP ratio at a summer mean TP concentration of <100–150 μg L^?1. This effect was strongest in shallow lakes. This implies potentially higher rates of zooplankton grazing and may be ascribed to the observed large changes in fish community structure and biomass with decreasing TP contribution. In 82% of the lakes for which data on fish are available, fish biomass declined with TP. The percentage of piscivores increased in 80% of those lakes and often a shift occurred towards dominance by fish species characteristic of less eutrophic waters. 6. Data on macrophytes were available only for a small subsample of lakes. In several of those lakes, abundance, coverage, plant volume inhabited or depth distribution of submerged macrophytes increased during oligotrophication, but in others no changes were observed despite greater water clarity. 7. Recovery of lakes after nutrient loading reduction may be confounded by concomitant environmental changes such as global warming. However, effects of global change are likely to run counter to reductions in nutrient loading rather than reinforcing re‐oligotrophication.     

Seasonal variation in phytoplankton composition and physical-chemical features of the shallow Lake Doïrani,Macedonia, Greece

Phytoplankton species composition, seasonal dynamics and spatial distribution in the shallow Lake Doïrani were studied during the growth season of 1996 along with key physical and chemical variables of the water. Weak thermal stratification developed in the lake during the warm period of 1996. The low N:P ratio suggests that nitrogen was the potential limiting nutrient of phytoplankton in the lake. In the phytoplankton of the lake, Chlorophyceae were the most species-rich group followed by Cyanophyceae. The monthly fluctuations of the total phytoplankton biomass presented high levels of summer algal biomass resembling that of other eutrophic lakes. Dinophyceae was the group most represented in the phytoplankton followed by Cyanophyceae. Diatomophyceae dominated in spring and autumn. Nanoplankton comprised around 90% of the total biomass in early spring and less than 10% in summer. The seasonal dynamics of phytoplankton generally followed the typical pattern outlined for other eutrophic lakes. R-species (small diatoms), dominant in the early phase of succession, were replaced by S-species (Microcystis, Anabaena, Ceratium) in summer. With cooling of the water in September, the biomass of diatoms (R-species) increased. The summer algal maxima consisted of a combination of H and M species associations (sensu Reynolds). Phytoplankton development in 1996 was subject to the combined effect of the thermal regime, the small depth of mixing and the increased sediment-water interactions in the lake, which caused changes in the underwater light conditions and nutrient concentrations.     

Light limitation of phytoplankton development in an oligotrophic lake - Loch Ness, Scotland

• 1 The underwater light climate in Loch Ness is described in terms of mixing depth (Z_m) and depth of the euphoric zone (Z_eu). During periods of complete mixing, Z_m equates with the mean depth of the loch (132 m), but even during summer stratification the morphometry of the loch and the strong prevailing winds produce a deep thermocline and an epilimnetic mixed layer of about 30 m or greater. Hence, throughout the year the quotient Z_m/Z_eu is exceptionally high and the underwater light climate particularly unfavourable for phytoplankton production and growth.
• 2 Phytoplankton biomass expressed as chlorophyll a is very low in Loch Ness, with a late summer maximum of less than 1.5 mg chlorophyll a m^-3 in the upper 30 m of the water column. This low biomass and the resulting very low photosynthetic carbon fixation within the water column are evidence that a severe restraint is imposed on the rate at which phytoplankton can grow in the loch.
• 3 The chlorophyll a content per unit of phytoplankton biovolume and the maximum, light-saturated specific rate of photosynthesis are both parameters which might be influenced by the light climate under which the phytoplankton have grown. However, values obtained from Loch Ness for both chlorophyll a content (mean 0.0045 mg mm^-3) and maximum photosynthetic rate (1–4 mg C mg Chla^-1 h^-1) are within the range reported from other lakes.
• 4 Laboratory bioassays with the natural phytoplankton community from Loch Ness on two occasions in late summer when the light climate in the loch is at its most favourable, suggest that even then limitation of phytoplankton growth is finely balanced between light and phosphorus limitation. Hence, for most of the year, when the light climate is less favourable, phytoplankton growth will be light limited.
• 5 Quotients relating mean annual algal biomass as chlorophyll a (c. 0.5 mg Chla m^-3) and the probable annual specific areal loading of total phosphorus (0.4–1.7 g TP m^-2 yr^-1) suggest that the efficiency with which phytoplankton is produced in Loch Ness per unit of TP loading is extremely low when compared with values from other Scottish lochs for which such an index has been calculated. This apparent inefficiency can be attributed to suppression of photosynthetic productivity in the water column due to the unfavourable underwater light climate.
• 6 These several independent sources of evidence lead to the conclusion that phytoplankton development in Loch Ness is constrained by light rather than by nutrients. Loch Ness thus appears to provide an exception to the generally accepted paradigm that phytoplankton development in lakes of an oligotrophic character is constrained by nutrient availability.

     

珠江口及毗邻海域营养盐对浮游植物生长的影响

基于2006年7月(夏季),10月(秋季)和2007年3月(春季)的现场调查数据,对珠江口及毗邻海域中的营养盐和叶绿素a等环境生态因子的时空分布特性进行了对比分析,研究了氮磷比与叶绿素a含量和种群多样性之间的联系,探讨了该海域营养盐对于浮游植物生长的影响。结果表明:(1)研究海域营养盐表现出较强的季节和空间差异性,总氮(TN)和总磷(TP)浓度均值春季(1.545 mg/L、0.056 mg/L)和夏季(1.570 mg/L、0.058 mg/L)均大于秋季(1.442 mg/L、0.034 mg/L),且春夏季浓度空间差异更明显。(2)调查期间海域营养盐含量超标现象突出,夏季尤为明显。无机氮(DIN)总体均值0.99 mg/L,超四类海水标准限值1倍,活性磷酸盐(PO4-P)总体均值0.021 mg/L,DIN∶PO4-P平均值为130;叶绿素a浓度与营养盐、p H、温度有较显著的相关性。(3)叶绿素a浓度较高的站位,具有较高的DIN∶PO4-P值,但浮游植物多样性指数偏低,优势种明显,主要为中肋骨条藻。氮磷比的改变会影响不同生长特性的浮游植物间的竞争和种群结构的改变;今后海洋污染治理中,在控制氮、磷污染时要注意氮磷比的改变可能造成的浮游生态影响。     

Does high nitrogen loading prevent clear‐water conditions in shallow lakes at moderately high phosphorus concentrations?

1. The effect of total nitrogen (TN) and phosphorus (TP) loading on trophic structure and water clarity was studied during summer in 24 field enclosures fixed in, and kept open to, the sediment in a shallow lake. The experiment involved a control treatment and five treatments to which nutrients were added: (i) high phosphorus, (ii) moderate nitrogen, (iii) high nitrogen, (iv) high phosphorus and moderate nitrogen and (v) high phosphorus and high nitrogen. To reduce zooplankton grazers, 1⁺ fish (Perca fluviatilis L.) were stocked in all enclosures at a density of 3.7 individuals m^?2. 2. With the addition of phosphorus, chlorophyll a and the total biovolume of phytoplankton rose significantly at moderate and high nitrogen. Cyanobacteria or chlorophytes dominated in all enclosures to which we added phosphorus as well as in the high nitrogen treatment, while cryptophytes dominated in the moderate nitrogen enclosures and the controls. 3. At the end of the experiment, the biomass of the submerged macrophytes Elodea canadensis and Potamogeton sp. was significantly lower in the dual treatments (TN, TP) than in single nutrient treatments and controls and the water clarity declined. The shift to a turbid state with low plant coverage occurred at TN >2 mg N L^?1 and TP >0.13–0.2 mg P L^?1. These results concur with a survey of Danish shallow lakes, showing that high macrophyte coverage occurred only when summer mean TN was below 2 mg N L^?1, irrespective of the concentration of TP, which ranged between 0.03 and 1.2 mg P L^?1. 4. Zooplankton biomass and the zooplankton : phytoplankton biomass ratio, and probably also the grazing pressure on phytoplankton, remained overall low in all treatments, reflecting the high fish abundance chosen for the experiment. We saw no response to nutrition addition in total zooplankton biomass, indicating that the loss of plants and a shift to the turbid state did not result from changes in zooplankton grazing. Shading by phytoplankton and periphyton was probably the key factor. 5. Nitrogen may play a far more important role than previously appreciated in the loss of submerged macrophytes at increased nutrient loading and for the delay in the re‐establishment of the nutrient loading reduction. We cannot yet specify, however, a threshold value for N that would cause a shift to a turbid state as it may vary with fish density and climatic conditions. However, the focus should be widened to use control of both N and P in the restoration of eutrophic shallow lakes.     

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 mixing and silica enrichment on phytoplankton seasonal succession

During the summer of 1983, cryptophytes, diatoms, cyanophytes, and the dinoflagellate, Ceratium hirundinella were most prominant among the phytoplankton of Eau Galle Reservoir. In the open water, cryptophytes and diatoms peaked in the spring, cyanophytes were most successful in the early summer, and Ceratium was dominant from mid-July until early August. In general, the sequence of events corresponded quite closely to the model of seasonal succession developed by the Plankton Ecology Group of the International Society of Limnology. To a large extent, the same pattern held in four experimental water columns. Departures from the model involved the roles of specific nutrients in diatom and cyanophyte periodicity. Diatoms began to yield to cyanophytes in late spring despite intermittent mixing and silica enrichment. Although capable of buoyancy regulation and thus well adapted to stable water columns, cyanophytes had greater increases in biomass in mixed columns, and in those columns, were most successful during a period of intermittent mixing. Cyanophyte success varied inversely with TN : TP ratios during the period of intermittent mixing, but not subsequently. By mid-July, Ceratium dominated the phytoplankton of every column except that of a mixed column in which conditions favored cyanophytes and large diatom species.     

Nutrient removal from polluted stream water by artificial aquatic food web system

For the removal of nutrients from eutrophic stream water polluted by non-point sources, an artificial aquatic food web (AAFW) system comprising processes of phytoplankton growth and Daphnia magna grazing was developed. The AAFW system was a continuous-flow system constructed with one storage basin of 3 m³ capacity, one phytoplankton tank of 3 m³ capacity, and one zooplankton growth chamber of 1.5 m³ capacity. The system was optimized by setting hydraulic retention time of phytoplankton tank as 3 days and D. magna density as 740–1000 individual l⁻¹. When the system was operated on eutrophic stream water that was delivering 471 g of total nitrogen (TN) and 29 g of total phosphorus (TP) loadings for 45 days, 250 g (53%) of TN and 16 g (54%) of TP were removed from the water during its passage through the phytoplankton tank. In addition, 64 g (14%) of TN and 4 g (13%) of TP were removed from the water by harvesting zooplankton biomass in the zooplankton growth chamber, resulting in significant overall removal rates of TN (69%), nitrate (78%), TP (73%), and dissolved inorganic phosphorus (94%). While the removal efficiency of the AAFW system is comparable to those of other ecotechnologies such as constructed wetlands, its operation is less limited by the availability of space or seasonal shift of temperature. Therefore, it was concluded that AAFW system is a highly efficient, flexible system for reducing nutrient levels in tributary streams and hence nutrient loading to large aquatic systems receiving the stream water. Handling editor: J. Padisak