Ambiguous climate impacts on competition between submerged macrophytes and phytoplankton in shallow lakes

1. Shallow lakes may switch from a state dominated by submerged macrophytes to a phytoplankton‐dominated state when a critical nutrient concentration is exceeded. We explore how climate change may affect this critical nutrient concentration by linking a graphical model to data from 83 lakes along a large climate gradient in South America. 2. The data indicate that in warmer climates, submerged macrophytes may tolerate more underwater shade than in cooler lakes. By contrast, the relationship between phytoplankton biomass [approximated by chlorophyll‐a (chl‐a) or biovolume] and nutrient concentrations did not change consistently along the climate gradient. In warmer climates, the correlation between phytoplankton biomass and nutrient concentrations was overall weak, especially at low total phosphorus (TP) concentrations where the chl‐a/ TP ratio could be either low or high. 3. Although the enhanced shade tolerance of submerged plants in warmer lakes might promote the stability of their dominance, the potentially high phytoplankton biomass at low nutrient concentrations suggests an overall low predictability of climate effects. 4. We found that near‐bottom oxygen concentrations are lower in warm lakes than in cooler lakes, implying that anoxic P release from eutrophic sediment in warm lakes likely causes higher TP concentrations in the water column. Subsequently, this may lead to a higher phytoplankton biomass in warmer lakes than in cooler lakes with similar external nutrient loadings. 5. Our results indicate that climate effects on the competitive balance between submerged macrophytes and phytoplankton are not straightforward.     

浅水湖泊浮水植物芡实收割对浮游植物群落的影响

浅水湖泊生态系统正遭受广泛而强烈的人为干扰,但是对收割水生植物干扰的研究甚少。于2019年8月对芡实过度生长的陈瑶湖进行通道式分区收割工程,分析了收割芡实（Euryale ferox）前后不同处理组浮游植物群落的变化。研究期间共鉴定出浮游植物6门47属72种,其中收割前63种,收割后71种。收割后浮游植物的细胞密度和生物量均高于收割前,分别增加了39.78%和5.09%。收割芡实导致陈瑶湖浮游植物群落为由蓝藻-绿藻-硅藻-隐藻群落转变为蓝藻-绿藻-硅藻群落。其中蓝藻细胞密度和生物量显著高于收割前（P<0.05）,归因于有害蓝藻（铜绿微囊藻Microcystis aeruginosa、水华束丝藻Aphanizomenon flos-aquae、小颤藻Oscillatoria tenuis、卷曲鱼腥藻Dolicospermum circinale、小席藻Phormidium tenu）的增加。收割还导致了硅藻群落由附生型向浮游型硅藻的转变,表现为尖针杆藻（Ulnaria acus）减少,而颗粒直链藻极狭变种（Aulacoseira granulata var.angustissima）、梅尼小环藻（Cyclotella meneghiniana）增加。在芡实收割过程中,未收割组和河道的浮游植物群落结构在时空分布上无显著性差异（P>0.05）,但收割组在收割后的不同阶段内差异较为明显,其细胞密度和生物量随着收割实验的结束逐渐降低。浮游植物与环境因子的相关性分析表明,水生植被覆盖度、总磷、总氮、溶解氧和叶绿素a浓度是影响浮游植物细胞密度和生物量变化的主要环境因子。综合陈瑶湖水质状态,本研究认为收割芡实并不能缓解浅水湖泊富营养化状况,研究结果为浅水湖泊水生植被的管理提供理论依据。     

Phytoplankton and submerged macrophytes in the aquatic ecosystems of the Danube Delta during the last decade

The dynamics are presented of the main primary producers in seven representative Danube Delta lakes. Generally, the primary productivity developed from the macrophyte-epiphyte complex towards the phytoplankton. Species changes occurred in the two main compartments with increasing relative abundance of colonial blue-green algae in the phytoplankton and in submerged macrophytes of species with a vertical growth strategy. These changes are linked to accelerated eutrophication of the lakes, with increased phosphorus loading and a reduction in N:P ratio.     

Aquatic macrophytes and phytoplankton of a quarry lake in the upper Vicenza range: preliminary study

Abstract

The work was carried out during 1995 in a quarry-lake and 13 samplings were performed to gather data on the aquatic flora. Sampling locations both along the shores and in the water provided data to define the position and the amount of covering by macrophytes. As regards the microphytes, considered as the sole phytoplankton component, the composition and the seasonal abundance were evaluated.     

Climate-related differences in the dominance of submerged macrophytes in shallow lakes

It has been suggested that shallow lakes in warm climates have a higher probability of being turbid, rather than macrophyte dominated, compared with lakes in cooler climates, but little field evidence exists to evaluate this hypothesis. We analyzed data from 782 lake years in different climate zones in North America, South America, and Europe. We tested if systematic differences exist in the relationship between the abundance of submerged macrophytes and environmental factors such as lake depth and nutrient levels. In the pooled dataset the proportion of lakes with substantial submerged macrophyte coverage (> 30% of the lake area) decreased in a sigmoidal way with increasing total phosphorus (TP) concentration, falling most steeply between 0.05 and 0.2 mg L⁻¹. Substantial submerged macrophyte coverage was also rare in lakes with total nitrogen (TN) concentrations above 1–2 mg L⁻¹, except for lakes with very low TP concentrations where macrophytes remain abundant until higher TN concentrations. The deviance reduction of logistic regression models predicting macrophyte coverage from nutrients and water depth was generally low, and notably lowest in tropical and subtropical regions (Brazil, Uruguay, and Florida), suggesting that macrophyte coverage was strongly influenced by other factors. The maximum TP concentration allowing substantial submerged macrophyte coverage was clearly higher in cold regions with more frost days. This is in agreement with other studies which found a large influence of ice cover duration on shallow lakes' ecology through partial fish kills that may improve light conditions for submerged macrophytes by cascading effects on periphyton and phytoplankton. Our findings suggest that, in regions where climatic warming is projected to lead to fewer frost days, macrophyte cover will decrease unless the nutrient levels are lowered.     

Typology of shallow lakes of the Salado River basin (Argentina), based on phytoplankton communities

The phytoplankton communities of eleven shallow lakes from Buenos Aires Province, Argentina, were studied seasonally from 1987 to 1989. Several physical and chemical properties were measured in each lake (pH, temperature, dissolved oxygen, transparency, nutrients), in order to interpret the structural and dynamic traits of the phytoplankton community.Important differences between the lakes studied were put in evidence by means of multivariate techniques (Cluster Analysis and PCA). The shallow lakes densely populated by macrophytes hosted lowest phytoplanktonic densities, with average values ranging from 690 to 16500 algae ml^–1. High species diversities were observed in these lakes (4.0–4.8). Lakes less colonized by macrophytes had higher phytoplankton densities. In some of them important blooms of Cyanophyceae were recorded, with between 60 000 and 179 000 algae ml^–1, and concomitant low diversities.The results of this investigation support the hypothesis that the phytoplankton community is strongly influenced by the macrophytes, by direct competition and/or by competition from periphytic algae associated with higher plants.     

An assessment of the importance of emergent and floating-leaved macrophytes to trophic status in the Loosdrecht lakes (The Netherlands)

The potential importance of the six major emergent and floating-leaved macrophyte species in recycling of sediment phosphorus in the Loosdrecht lakes was studied. Representative plant samples were collected at the time of maximum biomass, and analysed for biomass and carbon, nitrogen and phosphorus contents. Species cover was determined by aerial photography.Total cover in the seven lakes studied ranged between 2 and 26 percent. For the four main species, biomass per unit area increased with lake trophic status. Consistent differences in C, N and P contents per unit biomass were not observed. Although cover values were small, significant amounts of C, N and P were contained in the macrophytes when compared with maximum sestonic content.Potential P loads from macrophyte decay were calculated. In Lake Loosdrecht, the P load represented 15 percent of current external P inputs. The potential importance of macrophyte decay to P recycling in the other lakes is greater.Decay of macrophyte species at the end of the growing season appears to affect autumnal nutrient and chlorophyll a levels in the water column of some lakes. The re-establishment of submerged species following lake restoration may increase the importance of this pathway in the lakes.     

The role of periphytes in the shift between macrophyte and phytoplankton dominated systems in a shallow, eutrophic lake (Lake Taihu, China)

Based on experiments of periphyte response to different trophic levels and their impact on macrophyte production, it was found that the periphyte biomass increased with the nutrient concentrations. Increased trophic level and periphyte biomass resulted in decreased macrophyte photosynthesis. It was suggested that the periphytes could cause resilience and hysteresis in the system shifts between macrophyte and phytoplankton domination. Other factors, such as fish farming, storm induced waves and mechanical destruction, and high water levels could be the perturbations during the system shifts, but these are not the key factors. Instead, the nutrient loading and periphyte abundance could determine the shift in lake ecosystem between macrophyte and phytoplankton domination. This finding could theoretically elucidate the mechanism of ecosystem shifts between macrophyte and phytoplankton domination.     

Seasonal dynamics of phytoplankton and phytobenthos,and associated chemical interactions,in a shallow upland lake (Malham Tarn,northern England)

Seasonal changes of phytoplankton were followed over 3 years (1985–87) in a shallow, unstratified and calcareous upland lake.The phytoplankton was of low to moderate abundance and generally dominated by phytoflagellates. Seasonality involved a winter minimum of abundance, a spring maximum of diatoms, and often brief increases in summer that included blue-greens, especially the colonial Gloeotrichia echinulata. Some components were of benthic origin. Seasonal growth of the main component of the phytobenthos, Chara globularisvar. virgata, caused a regular summer depletion in lake water of Ca²⁺ and HCO₃ ^- (alkalinity) by associated CaCO₃ deposition, and a more extreme (and unusual) depletion of K⁺. Chemical analysis of Chara biomass and of underlying sediments indicated a large benthic nutrient stock, much surpassing that represented by the phytoplankton. Growth in this biomass, and the magnitude of water-borne inputs, influenced the removals of Ca²⁺, K⁺ and inorganic N. The phytoplankton was probably limited by a low-P medium, to which co-precipitation of phosphate with CaCO₃ may have contributed. A vernal depletion of Si was probably limiting to diatom growth, and appeared to be mainly induced by benthic rather than planktonic diatoms. Examples of long-term change in composition of the phytoplankton and phytobenthos are noted and discussed in relation to the interaction of these components, nutrient enrichment, and possible alternative stable states.     

The role of periphytes in the shift between macrophyte and phytoplankton dominated systems in a shallow, eutrophic lake (Lake Taihu, China)

Based on experiments of periphyte response to different trophic levels and their impact on macrophyte production,it was found that the periphyte biomass increased with the nutrient con-centrations. Increased trophic level and periphyte biomass resulted in decreased macrophyte photo-synthesis. It was suggested that the periphytes could cause resilience and hysteresis in the system shifts between macrophyte and phytoplankton domination. Other factors,such as fish farming,storm induced waves and mechanical destruction,and high water levels could be the perturbations during the system shifts,but these are not the key factors. Instead,the nutrient loading and periphyte abundance could determine the shift in lake ecosystem between macrophyte and phytoplankton domination. This finding could theoretically elucidate the mechanism of ecosystem shifts between macrophyte and phytoplankton domination.     

Weedbeds and big bugs: the importance of scale in detecting the influence of nutrients and predation on macroinvertebrates in plant‐dominated shallow lakes

1. The scale of investigations influences the interpretation of results. Here, we investigate the influence of fish and nutrients on biotic communities in shallow lakes, using studies at two different scales: (i) within‐lake experimental manipulation and (ii) comparative, among‐lake relationships. 2. At both scales, fish predation had an overriding influence on macroinvertebrates; fish reduced macroinvertebrate biomass and altered community composition. Prey selection appeared to be size based. Fish influenced zooplankton abundance and light penetration through the water column also, but there was no indication that fish caused increased resuspension of sediment. 3. There were effects of nutrients at both scales, but these effects differed with the scale of the investigation. Nutrients increased phytoplankton and periphyton at the within‐lake scale, and were associated with increased periphyton at the among‐lake scale. No significant effect of nutrients on macroinvertebrates was observed at the within‐lake scale. However, at the among‐lake scale, nutrients positively influenced the biomass and density of macroinvertebrates, and ameliorated the effect of fish on macroinvertebrates. 4. Increased prey availability at higher nutrient concentrations would be expected to cause changes in the fish community. However, at the among‐lake scale, differences were not apparent in fish biomass among lakes with different nutrient conditions, suggesting that stochastic events influence the fish community in these small and relatively isolated shallow lakes. 5. The intensity of predation by fish significantly influences macroinvertebrate community structure of shallow lakes, but nutrients also play a role. The scale of investigation influences the ability to detect the influence of nutrients on the different components of shallow lake communities, particularly for longer lived organisms such as macroinvertebrates, where the response takes longer to manifest.     

The effects of artificial enrichment upon the planktonic and benthic communities in a mesotrophic to hypertrophic loch series in lowland Scotland

Phytoplankton, submerged macrophytes, zoobenthos and zooplankton communities were studied concurrently in three lochs which progressively increased in trophic state. The phytoplankton showed a change in dominance from Bacillariophyceae (Pennales) and Desmidaceae in the mesotrophic Loch of the Lowes to Bacillariophyceae (Pennales) and Cyanobacteria in eutrophic Balgavies Loch and to Bacillariophyceae (Centrales) and Chlorophyceae in hypertrophic Forfar Loch. The Nygaard Compound phytoplankton quotient rose from 1.2 in the Lowes to 12 in Forfar. There was a decrease in number of species of submerged macrophytes, and in depth of colonisation from 5 m in the Lowes to 2 m in Forfar which was proportional to the decrease in mean transparency. Macrophyte biomass, however, increased in proportion to the concentration of total phosphorus in the lochs. There was a decrease in species of littoral zoobenthos, but an increase in density of profundal benthos from the Lowes to Forfar. Zooplankton also showed an increase in density with enrichment, accompanied by changes in dominance. Calanoida were dominant in the Lowes; Cladocera in Balgavies and Cyclopoda in Forfar. The importance of these differences in understanding the effects of progressive enrichment and to the management of the lochs is discussed.     

Managing water quality with aquatic macrophytes

The principal sources of water for human use are lakes, rivers, soil moisture and relatively shallow groundwater basins. Water quality in lakes and reservoirs is subjected to the natural degradation, processes of eutrophication and the impacts of human activities. Water quality problems can often be as severe as those of water availability but less attention has been paid to them, particularly in developing countries. Currently additional sustainable ways to mitigate the degradation of water quality are being researched all over the world. Phytoremediation is one of the serious efforts towards the sustainability. Most of the aquatic macrophytes are naturally occurring and well adapted for their surroundings. Aquatic macrophytes have the capability to remove excessive nutrient load from the water that otherwise cause eutrophication of the water body. Aquatic macrophytes absorb nutrient mineral ions from water column and influence metal retention indirectly by acting as traps for particulate matter, by slowing the water current and favoring sedimentation of suspended particles. Aquatic macrophytes also reduce sediment resuspension by offering wind protection. The use of aquatic macrophyte for treatment of wastewater to mitigate variety of pollution level is one of the most researched issues all over the world. Aquatic plant species are very specific for the uptake of nutrients. Owing to this specificity, the selection of the aquatic plant species is one of the skilled tasks prior to the design of a water treatment facility. An effort has been made in this review to cover the most researched aquatic flora for mitigation purposes and their possible use in a mesocosm as the selection of an appropriate aquatic plant specie reduce the time and cost of the treatment processes.     

Engineering and biological approaches to the restoration from eutrophication of shallow lakes in which aquatic plant communities are important components

Engineering approaches (nutrient removal, sediment pumping, hypolimnion oxygenation, alum treatments) may be most appropriate to deep lakes where the aim of restoration from eutrophication is simply to reduce the production and crop of one component, the phytoplankton. They do not always give the desired results because the nutrient loading may only be reduced to a limited extent. There are additional problems in shallow lakes where change of state between community dominance (aquatic plants versus plankton) is wanted. Each community has powerful buffering mechanisms and biomanipulation may be essential to switch one state to another even with considerable nutrient reduction. For the phytoplankton-dominated community the buffers include the advantages of early growth, lower diffusion pathways for CO₂, overhead shading, and an absence of large cladoceran grazers. This later is because open-water shallow environments provide no refuges against predation for the large Cladocera which are both the most efficient grazers and the most favoured prey for fish. Restoration of aquatic plants may then require provision of refuges for the grazers. Different sorts of refuge are discussed using case studies of Hoveton Great Broad and Cockshoot Broad in the Norfolk Broadland.     

Effects of common carp (Cyprinus carpio) on macrophytes and invertebrate communities in a shallow lake

1. We performed two field experiments using different cage sizes to determine the direct and indirect effects of carp on macrophytes and invertebrate community composition in a shallow lake. 2. The presence of carp produced a significant decrease in macrophyte abundance, changes in species composition and decreased abundance and diversity of invertebrates in small (2.5 × 2.5 × 2.0 m) cages. In large exclosures (15 × 5 m), only macrophyte stem length was significantly affected by carp. 3. There was considerable variation in the macrophyte and invertebrate responses to carp on different sides of the lake in the large exclosures, suggesting that wind and wave action are also important variables affecting macrophyte growth and persistence. 4. The data suggest that carp can significantly affect species abundance and diversity of macrophytes and some macroinvertebrates.     

Long-term compositional response of phytoplankton in a shallow,turbid environment,Neusiedlersee (Austria/Hungary)

Data on phytoplankton biomass and on nutrient concentrations from Neusiedler See (mean depth 1.3 m) covering more than two decades are presented. The lake underwent strong eutrophication during this period. The response of annual average phytoplankton biomass and chlorophyll-a to the increase of phosphorus concentration from 10 to > 100 µg l^-1 was moderate (7-fold increase). This is caused by light limitation of the system because of the high inorganic turbidity of the lake. Analyses of the spring, summer and autumn seasons at the generic and higher taxonomic levels show significant changes in composition of the phytoplankton community. Diatoms were more important during the pre-eutrophication phase while Chlorophyceae became most prominent during the peak of the eutrophication process. Blue-green algae, including Microcystis, became more apparent after this period. The abundance of some groups or genera, e.g. Euglena, was linked to the decline and re-appearance of submerged macrophytes in the lake. Abiotic and biotic interactions as causes for the observed changes are discussed.     

Biological control of phytoplankton by the subtropical submerged macrophytes Egeria densa and Potamogeton illinoensis: a mesocosm study

1. In temperate regions, submerged macrophytes can hamper phytoplankton blooms. Such an effect could arise directly, for instance via allelopathy, or indirectly, via competition for nutrients or the positive interaction between submerged macrophytes and zooplankton grazing. However, there is some evidence that the positive interaction between submerged macrophytes and zooplankton grazing is less marked in warmer regions, where the interaction is less well studied, and that negative effects of higher water plants on phytoplankton biomass are weaker. 2. We carried out two consecutive mesocosm experiments in Uruguay (subtropical South America) to study the effects of two common submerged macrophytes from this region (Egeria densa and Potamogeton illinoensis) on phytoplankton biomass, in the absence of zooplankton grazing. We compared phytoplankton development between different macrophyte treatments (no macrophytes, artificial macrophytes, real Egeria and real Potamogeton). We used artificial macrophytes to differentiate between physical effects (i.e. shading, sedimentation and competition with periphyton) and biological effects (i.e. nutrient competition and allelopathy). 3. In Experiment 1, we found no evidence for physical effects of macrophytes on phytoplankton biomass, but both macrophyte species seemed to exert strong biological effects on phytoplankton biomass. Only Egeria affected phytoplankton community structure, particularly tempering the dominance of Scenedesmus. Nutrient addition assays revealed that only Egeria suppressed phytoplankton through nutrient competition. 4. We performed a second mesocosm experiment with the same design, but applying saturating nutrient conditions as a way of excluding the effects of competition for nutrients. This experiment showed that both macrophytes were still able to suppress phytoplankton through biological mechanisms, providing evidence for allelopathic effects. Our results indicate that both common macrophytes are able to keep phytoplankton biomass low, even in the absence of zooplankton grazing.     

Clear,crashing, turbid and back – long‐term changes in macrophyte assemblages in a shallow lake

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