Lake‐type‐specific seasonal patterns of nutrient limitation in German lakes,with target nitrogen and phosphorus concentrations for good ecological status

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Stoichiometric relationships in vernal pond plankton communities

1. The light-nutrient hypothesis (LNH) predicts that changes in light supply can alter the balance of nutrient and energy limitation in primary producers. We tested this prediction by examining temporal changes in vernal forest ponds, which are highly dynamic systems with respect to seasonal change in light and nutrient supply. In three vernal ponds that differ in productivity, we measured changes in light, total and seston nitrogen and phosphorus, and seston carbon and chlorophyll during the spring, before and after tree leaf-out. We also quantified changes in the population dynamics of the major zooplankton grazers in these systems.
2. In each pond, nutrient levels increased and light levels declined, creating a temporal shift in light-nutrient supply to the plankton. Results generally supported predictions of stoichiometric theory and the LNH, but there were notable exceptions.
3. Seston C : N : P ratios rapidly changed in response to dramatic increases in N and P supply rates. However, seston N : P was typically lower than values for total N : P in the water. Furthermore, as predicted, we observed a decline in seston C : P as the light : nutrient ratio declined, but seston C : N simultaneously increased. These results suggest an unexpected shift towards potential nitrogen limitation. Alternatively, this change in nutrient ratios may be driven by a seasonal change in phytoplankton composition or nutritional mode.
4. Seston carbon concentrations remained stable despite seasonal changes in grazing intensity associated with the phenology of large-bodied Daphnia grazers. However, chlorophyll concentrations declined dramatically as the season progressed, resulting in a simultaneous decline in the C : Chlorophyll ratio of seston. Both pond shading and increased grazing probably contributed to the decline in chlorophyll.     

Concentrations of nitrogen and phosphorus in streams draining the mixed land-use Dee Catchment, north-east Scotland

1.  There is still some uncertainty about large-scale influences on nutrient budgets in rivers. In particular, reduced forms of nitrogen (N) in organic forms might represent a significant fraction of the soluble N present in headwater streams, but this is not well quantified. Nitrate increases in relative importance downstream within agriculturally dominated areas. Here we appraise variations in N dynamics for a representative temperate but upland river, the Dee.
2.  In the Dee catchment, the source of organic N appears to vary seasonally. During summer under low flow conditions it originates primarily from in-stream biological production, while during the winter–spring period leaching from the plant–soil system would be the major contributor.
3.  On any individual sampling day, a wide range of N : P ratios can occur in the catchment area. Generally the narrowest N : P ratios occur during the summer and early autumn, particularly for upland catchments dominated by semi-natural vegetation. It is possible that some of the tributaries and upper region of the main river may be limited by N during the summer. The interpretation of the N : P ratios depends greatly upon the potential biological availability of the organic N, which remains unknown.
4.  Together, these data further illustrate that simple ideas about the relative limiting effects of N and P in temperate freshwaters may be misleading.     

Nutrient dynamics and successional changes in a lentic freshwater biofilm

SUMMARY 1. Colonisation, species composition, succession of microalgae and nutrient dynamics in biofilms grown under light and dark conditions were examined during the initial phases of biofilm development in a lentic freshwater environment.
2. Biofilms were developed on inert (perspex) panels under natural illuminated and experimental dark conditions and the panels were retrieved for analysis after different incubation periods. Analysed parameters included biofilm thickness, algal density, biomass, chlorophyll a , species composition, total bacterial density and nutrients such as nitrite, nitrate, phosphate and silicate.
3. Biofilm thickness, algal density, biomass, chlorophyll a and species richness were significantly higher in light-grown biofilms, compared with dark-grown biofilms. The light-grown biofilms showed a three-phased succession pattern, with an initial domination of Chlorophyceae followed by diatoms (Bacillariophyceae) and finally by cyanobacteria. Dark-grown biofilms were mostly dominated by diatoms.
4. Nutrients were invariably more concentrated in biofilms than in ambient water. Nutrient concentrations were generally higher in dark-grown biofilms except in the case of phosphate, which was more concentrated in light-grown biofilms. Significant correlations between nutrients and biofilm parameters were observed only in light-grown biofilms.
5. The N : P ratio in the biofilm matrix decreased sharply in the initial 4 days of biofilm growth; ensuing N-limitation status seemed to influence biofilm community structure. The N : P ratios showed significant positive correlations with the chlorophycean fraction in both light and dark-grown biofilms, and low N : P ratio in the older biofilms favoured cyanobacteria. Our data indicate that nutrient chemistry of biofilm matrix shapes community structure in microalgal biofilms.     

Inverse relationship of epilithic algae and pelagic phosphorus in unproductive lakes: Roles of N2 fixers and light

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The role of light for fish–zooplankton–phytoplankton interactions during winter in shallow lakes – a climate change perspective

1. Variations in the light regime can affect the availability and quality of food for zooplankton grazers as well as their exposure to fish predation. In northern lakes light is particularly low in winter and, with increasing warming, the northern limit of some present-day plankton communities may move further north and the plankton will thus receive less winter light.
2. We followed the changes in the biomass and community structure of zooplankton and phytoplankton in a clear and a turbid shallow lake during winter (November–March) in enclosures both with and without fish and with four different light treatments (100%, 55%, 7% and <1% of incoming light).
3. In both lakes total zooplankton biomass and chlorophyll- a were influenced by light availability and the presence of fish. Presence of fish irrespective of the light level led to low crustacean biomass, high rotifer biomass and changes in the life history of copepods. The strength of the fish effect on zooplankton biomass diminished with declining light and the effect of light was strongest in the presence of fish.
4. When fish were present, reduced light led to a shift from rotifers to calanoid copepods in the clear lake and from rotifers to cyclopoid copepods in the turbid lake. Light affected the phytoplankton biomass and, to a lesser extent, the phytoplankton community composition and size. However, the fish effect on phytoplankton was overall weak.
5. Our results from typical Danish shallow eutrophic lakes suggest that major changes in winter light conditions are needed in order to have a significant effect on the plankton community. The change in light occurring when such plankton communities move northwards in response to global warming will mostly be of modest importance for this lake type, at least for the rest of this century in an IPCC A2 scenario, while stronger effects may be observed in deep lakes.     

Nutrient limitation of bacterioplankton and phytoplankton in humic lakes in northern Sweden

1. Two small humic lakes in northern Sweden with concentrations of dissolved organic carbon (DOC) between 15 and 20 mg L^–1 were fertilized with inorganic phosphorus (P) and inorganic nitrogen (N), respectively. A third lake was unfertilized and served as a control. In addition to this lake fertilization experiment, data from different regional surveys were used to assess the role of different limiting factors.
2. The P fertilization had no effects on bacterioplankton or phytoplankton, while phytoplankton were significantly stimulated by N fertilization. Inorganic nutrient limitation of bacterioplankton was a function of DOC concentration in water of the investigated region and nutrient-limited bacteria were found only in lakes with DOC concentrations less than around 15 mg L^–1
3. The fertilization experiments demonstrated that the DOC-rich experimental lakes contained a bioavailable pool of P that was not utilized to its full potential under natural conditions. The overall mobilization of energy (bacterioplankton plus phytoplankton) in the experimental lakes was restricted by lack of inorganic N.     

Modulation of the bacterial response to spectral solar radiation by algae and limiting nutrients

SUMMARY 1. The response of bacterial production (measured as [³H]TdR incorporation rate) to spectral solar radiation was quantified experimentally in an oligotrophic high-mountain lake over 2 years. Bacterial responses were consistent: ultraviolet-B (UVB) was harmful, whereas ultraviolet-A (UVA) + photosynthetically active radiation (PAR) and PAR enhanced bacterial activity. Full sunlight exerted a net stimulatory effect on bacterial activity in mid-summer but a net inhibitory effect towards the end of the ice-free period.
2. Experiments were undertaken to examine whether the bacterial response pattern depended on the presence of algae and/or was modulated by the availability of a limiting inorganic nutrient (phosphorus, P). In the absence of algae, [³H]TdR incorporation rates were significantly lower than when algae were present under all light treatments, and the consistent bacterial response was lost. This suggests that the bacterial response to spectral solar radiation depends on fresh-C released from algae, which determines the net stimulatory outcome of damage and repair in mid-summer.
3. In the absence of algae, UVB radiation inhibited bacteria when they were strongly P-deficient (mean values of N : P ratio: 46.1), whereas it exerted no direct effect on bacterial activity when they were not P-limited.
4. P-enrichment of lake water markedly altered the response of bacteria to spectral solar radiation at the end of ice-free period, when bacteria were strongly P-deficient. Phosphorus enrichment suppressed the inhibitory effect of full sunlight that was observed in October, both in whole lake water (i.e. including algae) and in the absence of algae. This indicates that the bacterial P-deficiency, measured as the cellular N : P ratio, was partly responsible for the net inhibitory effect of full sunlight, implying a high bacterial vulnerability to UVB.     

Global changes may be promoting a rise in select cyanobacteria in nutrient‐poor northern lakes

The interacting effects of global changes—including increased temperature, altered precipitation, reduced acidification and increased dissolved organic matter loads to lakes—are anticipated to create favourable environmental conditions for cyanobacteria in northern lakes. However, responses of cyanobacteria to these global changes are complex, if not contradictory. We hypothesized that absolute and relative biovolumes of cyanobacteria (both total and specific genera) are increasing in Swedish nutrient‐poor lakes and that these increases are associated with global changes. We tested these hypotheses using data from 28 nutrient‐poor Swedish lakes over 16 years (1998–2013). Increases in cyanobacteria relative biovolume were identified in 21% of the study sites, primarily in the southeastern region of Sweden, and were composed mostly of increases from three specific genera: Merismopedia, Chroococcus and Dolichospermum. Taxon‐specific changes were related to different environmental stressors; that is, increased surface water temperature favoured higher Merismopedia relative biovolume in low pH lakes with high nitrogen to phosphorus ratios, whereas acidification recovery was statistically related to increased relative biovolumes of Chroococcus and Dolichospermum. In addition, enhanced dissolved organic matter loads were identified as potential determinants of Chroococcus suppression and Dolichospermum promotion. Our findings highlight that specific genera of cyanobacteria benefit from different environmental changes. Our ability to predict the risk of cyanobacteria prevalence requires consideration of the environmental condition of a lake and the sensitivities of the cyanobacteria genera within the lake. Regional patterns may emerge due to spatial autocorrelations within and among lake history, rates and direction of environmental change and the niche space occupied by specific cyanobacteria.     

Primary production and phytoplankton composition in relation to DOC input and bacterioplankton production in humic Lake Örträsket

1. The biomass and production of picophytoplankton, large phytoplankton and heterotrophic bacterioplankton were measured in humic Lake Örträsket, northern Sweden during four consecutive summers.
2. High flow episodes, carrying fresh dissolved organic carbon (DOC) into the lake, always stimulated heterotrophic bacterial production at the expense of primary production. Primary production never exceeded bacterial production for approximately 20 days after such an episode had replenished epilimnial DOC. We suggest that allochthonous DOC is an energy source that stimulates bacterioplankton that, because of their efficient uptake of inorganic nutrients, are then able to outcompete phytoplankton. After the exhaustion of readily available DOC, phytoplankton were able to dominate epilimnion production in Lake Örträsket.
3. Biomass production was higher when dominated by phytoplankton than by bacterioplankton, despite a similar utilization of nutrients in the epilimnion throughout the summer. We propose that different C : N : P ratios of bacterioplankton and phytoplankton permit the latter to produce more carbon (C) biomass per unit of available inorganic nutrients than bacterioplankton.     

Geographical patterns of species turnover in aquatic plant communities

1. A classic theory in biogeography predicts that high latitude communities are unstable. This may be because of decreased species richness or decreased environmental predictability and productivity towards the poles.
2. We studied latitudinal patterns in long-term community persistence of aquatic vascular plants in 112 Finnish lakes, situated within a 1000-km range from the northernmost to the southernmost lake.
3. Contrary to theoretical predictions, we found that the turnover rate of plant species in 45 years was inversely related to latitude. That is, plant communities in northern lakes were more persistent than communities in southern lakes. When we used multiple regression to find the best predictors of species turnover rate (TR), latitude was the only variable that was highly significantly related to species turnover rate. Area, species number, water transparency, pH and change in transparency did not notably explain the gradient observed.
4. The latitudinal trend was mainly because of lower species immigration rates at higher latitudes, whereas extinction rate did not so strongly decrease with increasing latitude. Immigrations and extinctions in the lakes were not in balance: the species numbers between the 1930s and 1980s increased more strongly in the southern than northern lakes.
5. We suggest that the inverse relationship between latitude and plant species TR in Finland is most probably caused by human influence on lakes, especially eutrophication and immigration of new species in southern latitudes. In addition, although species richness per lake did not decrease towards the north, the total species pool probably does, which means that in the north there are fewer species that can actually immigrate.     

Nitrogen deposition and warming – effects on phytoplankton nutrient limitation in subarctic lakes

The aim of this study was to predict the combined effects of enhanced nitrogen (N) deposition and warming on phytoplankton development in high latitude and mountain lakes. Consequently, we assessed, in a series of enclosure experiments, how lake water nutrient stoichiometry and phytoplankton nutrient limitation varied over the growing season in 11 lakes situated along an altitudinal/climate gradient with low N‐deposition (<1 kg N ha^?1 yr^?1) in northern subarctic Sweden. Short‐term bioassay experiments with N‐ and P‐additions revealed that phytoplankton in high‐alpine lakes were more prone to P‐limitation, and with decreasing altitude became increasingly N‐ and NP‐colimited. Nutrient limitation was additionally most obvious in midsummer. There was also a strong positive correlation between phytoplankton growth and water temperature in the bioassays. Although excess nutrients were available in spring and autumn, on these occasions growth was likely constrained by low water temperatures. These results imply that enhanced N‐deposition over the Swedish mountain areas will, with the exception of high‐alpine lakes, enhance biomass and drive phytoplankton from N‐ to P‐limitation. However, if not accompanied by warming, N‐input from deposition will stimulate limited phytoplankton growth due to low water temperatures during large parts of the growing season. Direct effects of warming, allowing increased metabolic rates and an extension of the growing season, seem equally crucial to synergistically enhance phytoplankton development in these lakes.     

Thermal regimes of Florida lakes

Water column temperatures were determined monthly for 24 lakes and bimonthly for 5 lakes in peninsular Florida during 1979. Three geographical groups (north, central, south) were delineated from mean monthly water column temperatures for individual lakes. On a monthly basis, northern lakes were least similar to southern lakes, while central Florida lakes displayed greater affinity to the southern than to the northern lake group. Temperature differences between lake groups broke down during late summer. Subtropical lakes have been defined tentatively as those Florida lakes south of 28° latitude which possess warm monomictic circulation and a mean annual temperature of 24.2 ± 4.8 °C with minimum water column temperature rarely less than 14 °C and summer maxima rarely exceeding 31 °C. While all lakes in Florida are clearly warm monomictic annual nutrient cycling and productivity patterns may be influenced by inter-group differences in the timing and duration of water column circulation.     

A regional study of Holocene climate change and human occupation in Peruvian Amazonia

Aim  To investigate the influence of Holocene climatic and human-induced changes on a region of high biodiversity in southern Peruvian Amazonia.
Location  Four palaeoecological records from separate lakes within a lake district close to the modern city of Puerto Maldonado, Peru.
Results  The lakes provide a palaeoecological record spanning the last 8200 years. A mid-Holocene dry event is documented in all of the records that extend back > 6000 years. The dry event appears to have lasted from c . 7200 yr bp until c . 3300 yr bp . The onset of wetter conditions coincides with the formation of the youngest of the four lakes. The earliest occupation of these sites is inferred from the presence of charcoal at 7200 yr bp , and the first crop pollen is found at 3630 yr bp . Lakes that were regularly occupied were colonized soon after they formed. A reduction in charcoal concentration and the absence of crop pollen after c . 500  bp in all lakes is consistent with site abandonment following conquest.
Main conclusions  The mid-Holocene dry event is suggested to be part of a time-transgressive drying that tracked from north to south in both the Andes and the Amazon lowlands. The last millennium may represent the period of highest sustained lake levels within the Holocene. The proximity of the four lakes allows a landscape-scale analysis of the spatial extent of human disturbance centred on a known site of human occupation and reveals the highly localized nature of pre-Columbian anthropogenic disturbance in Amazonian landscapes. Inferences regarding widespread pre-Columbian landscape modification by indigenous peoples must take into account key site attributes, such as seasonality and proximity to rivers.     

Light and nutrient control phytoplankton biomass responses to global change in northern lakes

Global change affects terrestrial loadings of colored dissolved organic carbon (DOC) and nutrients to northern lakes. Still, little is known about how phytoplankton respond to changes in light and nutrient availability across gradients in lake DOC. In this study, we used results from whole‐lake studies in northern Sweden to show that annual mean phytoplankton biomass expressed unimodal curved relationships across lake DOC gradients, peaking at threshold DOC levels of around 11 mg/L. Whole‐lake single nutrient enrichment in selected lakes caused elevated biomass, with most pronounced effect at the threshold DOC level. These patterns give support to the suggested dual control by DOC on phytoplankton via nutrient (positively) and light (negatively) availability and imply that the lakes' location along the DOC axis is critical in determining to what extent phytoplankton respond to changes in DOC and/or nutrient loadings. By using data from the large Swedish Lake Monitoring Survey, we further estimated that 80% of northern Swedish lakes are below the DOC threshold, potentially experiencing increased phytoplankton biomass with browning alone, and/or combined with nutrient enrichment. The results support the previous model results on effects of browning and eutrophication on lake phytoplankton, and provide important understanding of how northern lakes may respond to future global changes.     

Drivers of long‐term invertebrate community stability in changing Swedish lakes

Research on ecosystem stability has had a strong focus on local systems. However, environmental change often occurs slowly at broad spatial scales, which requires regional‐level assessments of long‐term stability. In this study, we assess the stability of macroinvertebrate communities across 105 lakes in the Swedish “lakescape.” Using a hierarchical mixed‐model approach, we first evaluate the environmental pressures affecting invertebrate communities in two ecoregions (north, south) using a 23 year time series (1995–2017) and then examine how a set of environmental and physical variables affect the stability of these communities. Results show that lake latitude, size, total phosphorus and alkalinity affect community composition in northern and southern lakes. We find that lake stability is affected by species richness and lake size in both ecoregions and alkalinity and total phosphorus in northern lakes. There is large heterogeneity in the patterns of community stability of individual lakes, but relationships between that stability and environmental drivers begin to emerge when the lakescape, composed of many discrete lakes, is the focal unit of study. The results of this study highlight that broad‐scale comparisons in combination with long time series are essential to understand the effects of environmental change on the stability of lake communities in space and time.     

Browning reduces the availability—but not the transfer—of essential fatty acids in temperate lakes

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The response of a freshwater wetland to long-term ’low level‘ nutrient loads: nutrients and water budget

Boney Marsh is a small constructed freshwater wetland located along thefloodplain of the Kissimmee River in south Florida, USA. River water, withaverage Tot-P concentrations of 0.052 mg l⁻¹, Tot-N of 1.70 mgl⁻¹, and Cl⁻ of 15.95 mg l⁻¹, wasdiverted through the marsh to quantify mass retention and fate.Comprehensive mass balance budgets for Tot-P, Tot-N, and Cl⁻were developed based on input (inflow, precipitation) and output (outflow,evapotranspiration, seepage). Cl⁻, as well asNa⁺, budgets indicated that groundwater accounted forapproximately 7% of the total water budget. Annual mass loadings toBoney Marsh were 0.5, 15.7, and 147.9 g m⁻²year⁻¹ for Tot-P, Tot-N, and Cl⁻, respectively.Mean annual nutrient removal was estimated at 72% for Tot-P and34% for Tot-N, and P-assimilation capacity remained high andunchanged for the period of record. The subtropical marsh system accumulatedTot-P at a much higher rate than Tot-N, with averaged net sedimentationrates of 20.4 and 8.3 year⁻¹, respectively. Boney Marsh netsedimentation coefficients were higher than lakes with similar depths. TheN:P mass ratio in the wetland water column increased during the period ofrecord, and was primarily due to a high P-sedimentation rate and a decliningN-sedimentation rate. The authors use the editor‘s suggestion forabbreviations. This revised version was published online in July 2006 with corrections to the Cover Date.     

Effects of experimentally induced cyanobacterial blooms on crustacean zooplankton communities

SUMMARY 1. Large in situ enclosures were used to study the effects of experimentally induced cyanobacterial blooms on zooplankton communities. A combination of N and P was added to shallow (2 m) and deep enclosures (5 m) with the goal of reducing the TN : TP ratio to a low level (∼5 : 1) to promote cyanobacterial growth. After nutrient additions, high biomass of cyanobacteria developed rapidly in shallow enclosures reaching levels only observed during bloom events in eutrophic lakes.
2. In the shallow enclosures, particulate phosphorus (PP) was on average 35% higher in comparison with deep enclosures, suggesting that depth plays a key role in P uptake by algae. Phytoplankton communities in both deep and shallow enclosures were dominated by three cyanobacteria species – Aphanizomenon flos-aquae , Anabaena flos-aquae and Microcystis aeruginosa – which accounted for up to 70% of total phytoplankton biomass. However, the absolute biomass of the three species was much higher in shallow enclosures, especially Aphanizomenon flos-aquae . The three cyanobacteria species responded in contrasting ways to nutrient manipulation because of their different physiology.
3. Standardised concentrations of the hepatotoxic microcystin-LR increased as a result of nutrient manipulations by a factor of four in the treated enclosures. Increased biomass of inedible and toxin producing cyanobacteria was associated with a decline in Daphnia pulicaria biomass caused by a reduction in the number of individuals with a body length of >1 mm. Zooplankton biomass did not decline at moderate cyanobacteria biomass, but when cyanobacteria reached high biomass large cladocerans were reduced.
4. Our results demonstrate that zooplankton communities can be negatively affected by cyanobacterial blooms and therefore the potential to use herbivory to reduce algal blooms in such eutrophic lakes appears limited.     

Short-term and long-term effects of zooplanktivorous fish removal in a shallow lake: a synthesis of 15 years of data from Lake Zwemlust

SUMMARY 1. Removal of zooplanktivorous fish (mainly bream) in 1987 from a shallow eutrophic lake in the Netherlands, Lake Zwemlust, resulted in a quick switch from a turbid state with cyanobacteria blooms to a clear state dominated by macrophytes.
2. The clear state was not stable in the long term, however, because of high nutrient loadings.
3. In 1999, another removal of zooplanktivorous fish (mainly rudd) had similar effects as in 1987, although macrophytes returned more slowly.
4. In the years directly following both interventions there was a 'transition period' of very clear water with high densities of zooplanktonic grazers in the absence of macrophytes; low oxygen concentrations indicate that during those years primary production was low relative to heterotrophic activity.
5. The transition period appears to provide the light climate necessary for the return of macrophytes.
6. Reduction of nutrient loading is necessary to improve water quality in Lake Zwemlust in the long term. In the short term, repeated fish stock reduction is a reasonable management strategy to keep Lake Zwemlust clear.