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Reply to Cao et al.'s comment on “Does the responses of Vallisneria natans (Lour.) Hara to high nitrogen loading differ between the summer high-growth season and the lowgrowth season? Science of the Total Environment 601–602 (2017) 1513–1521”

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... Mesocosm enclosure is a suitable method to simulate how changes in temperature, nutrients and other environmental factors affect the long-term dynamics of aquatic organisms in shallow lake ecosystems [32,33]. To date, this method has been widely used to study the responses of phytoplankton [34], zooplankton [35], zoobenthos [36], and aquatic plants [37] to environmental change. ...
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To explore the impacts of multiple environmental stressors on animal communities in aquatic ecosystems, we selected protozoa—a highly sensitive group of organisms—to assess the effect of environmental change. To conduct this simulation we conducted a three-factor, outdoor, mesocosm experiment from March to November 2021. Changes in the community structure and functional group composition of protozoan communities under the separate and combined effects of these three environmental stressors were investigated by warming and the addition of nitrogen, phosphorus, and pesticides. The results were as follows: (1) Both eutrophication and pesticides had a considerable promotional effect on the abundance and biomass of protozoa; the effect of warming was not considerable. When warming was combined with eutrophication and pesticides, there was a synergistic effect and antagonistic effect, respectively. (2) Eutrophication promoted α diversity of protozoa and affected their species richness and dominant species composition; the combination of warming and pesticides remarkably reduced the α diversity of protozoa. (3) Warming, eutrophication, and pesticides were important factors affecting the functional groups of protozoa. Interaction among different environmental factors could complicate changes in the aquatic ecological environment and its protozoan communities. Indeed, in the context of climate change, it might be more difficult to predict future trends in the protozoan community. Therefore, our results provide a scientific basis for the protection and restoration of shallow lake ecosystems; they also offer valuable insights in predicting changes in shallow lakes.
... In turn, the morphologies and concentrations of C and N can directly affect the uptake of nutrients by primary productivity. For instance, a suitable ammonia (NH 4 + ) concentration is beneficial to the growth of phytoplankton, but excess NH 4 + will destroy the chloroplast ultrastructure and carbon skeleton [9,10]. In addition, the decomposition capacities of primary producers are also related to C, N and C/N ratios [11]. ...
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The metabolic activities of primary producers play an important role in the migration and transformation of carbon (C) and nitrogen (N) in aquatic environments. This study selected two typical areas in Taihu Lake, a cyanobacteria-dominant area (Meiliang Bay) and a macrophyte-dominant area (in the east area of the lake), to study the effects of cyanobacteria and macrophyte activities on C and N migration and transformation in aquatic environments. The results showed that total N and total particulate N concentrations in the water of the cyanobacteria-dominant area were much higher than those in the macrophyte-dominant area, which was mainly due to the assimilated intracellular N in cyanobacteria. Macrophyte activity drove a significantly higher release of dissolved organic C (DOC) in the water than that driven by cyanobacteria activity, and the DOC contents in the water of the macrophyte-dominant area were 2.4~4.6 times the DOC contents in the cyanobacteria-dominant area. In terms of the sediments, organic matter (OM), sediment total N and N species had positive correlations and their contents were higher in the macrophyte-dominant area than in the cyanobacteria-dominant area. Sediment OM contents in the macrophyte-dominant area increased from 4.19% to 9.33% as the sediment deepened (0~10 cm), while the opposite trend was presented in the sediments of the cyanobacteria-dominant area. Sediment OM in the macrophyte-dominant area may contain a relatively high proportion of recalcitrant OC species, while sediment OM in the cyanobacteria-dominant area may contain a relatively high proportion of labile OC species. Compared with the macrophyte-dominant area, there was a relatively high richness and diversity observed in the bacterial community in the sediments in the cyanobacteria-dominant area, which may be related to the high proportion of labile OC in the OM composition in its sediments. The relative abundances of most OC-decomposing bacteria, denitrifying bacteria, Nitrosomonas and Nitrospira were higher in the sediments of the cyanobacteria-dominant area than in the macrophyte-dominant area. These bacteria in the sediments of the cyanobacteria-dominant area potentially accelerated the migration and transformation of C and N, which may supply nutrients to overlying water for the demands of cyanobacteria growth. This study enhances the understanding of the migration and transformation of C and N and the potential effects of bacterial community structures under the different primary producer habitats.
... The amount of precipitation may disturb pond conditions, cause resuspension of the sediment, and create shading effects. When phytoplankton abundance increases, so do the levels of detritus and inorganic suspended matter (see Yu et al., 2018). Although the precipitation in this study was much higher than that in , the average Chla Phyt concentration (41.7 µg L −1 ) did not differ significantly from those in Yu et al. (2017) (33.8 µg L −1 ), indicating that the resuspension of the sediment caused by precipitation might not have had a significant effect on the macrophytes. ...
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Due to excess nutrient loading, loss of submersed macrophytes is a worldwide phenomenon in shallow lakes. Phosphorus is known to contribute significantly to macrophyte recession, but the role of nitrogen has received increasing attention. Our understanding of how high nitrogen concentrations affect the growth of submersed macrophytes, particularly under natural conditions, is still limited. In this study, we conducted experiments with canopy-forming Potamogeton crispus in 10 ponds subjected to substantial differences in nitrogen loading (five targeted total nitrogen concentrations: control, 2, 10, 20, and 100 mg L⁻¹) and compared the results with those of our earlier published experiments with rosette-forming Vallisneria natans performed 1 year before. Canopy-forming P. crispus was more tolerant than rosette-forming V. natans to exposure to high NH4 concentrations. This is probably because canopy-forming species reach the water surface where there is sufficient light for production of carbohydrates, thereby allowing the plants to partly overcome high NH4 stress. Both the canopy-forming P. crispus and the rosette-forming V. natans showed clear declining trends with increasing chlorophyll a in the water. Accordingly, shading by phytoplankton might be of key importance for the decline in submersed macrophytes in this experiment. Both experiments revealed free amino acids (FAA) to be a useful indicator of physiological stress by high ammonium but is not a reliable indicator of macrophyte growth.
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The global climate change may lead to more extreme climate events such as severe flooding creating excessive pulse-loading of nutrients, including nitrogen (N), to freshwaters. We conducted a 3-month mesocosm study to investigate the responses of phytoplankton, zooplankton and Vallisneria spinulosa to different N loading patterns using weekly and monthly additions of in total 14 g N m−2 month−1 during the first 2 months. The monthly additions led to higher phytoplankton chlorophyll a and total phytoplankton biomass than at ambient conditions as well as lower leaf biomass and a smaller ramet number of V. spinulosa. Moreover, the biomass of cyanobacteria was higher during summer (August) in the monthly treatments than those with weekly or no additions. However, the biomass of plankton and macrophytes did not differ among the N treatments at the end of the experiment, 1 month after the termination of N addition. We conclude that by stimulating the growth of phytoplankton (cyanobacteria) and reducing the growth of submerged macrophytes, short-term extreme N loading may have significant effects on shallow nutrient-rich lakes and that the lakes may show fast recovery if they are not close to the threshold of a regime shift from a clear to a turbid state.
Article
Yu et al.'s paper showed very interesting effects of high nitrogen (N) on the submerged macrophytes Vallisneria natans: active growth in the growing season enabled the macrophytes partly to overcome the ammonium stress. This result was evident in an experiment using ten pond ecosystems; however, their conclusion that shading induced by high phytoplankton biomass together with the toxicity of high ammonium contributed to the decrease of macrophytes growth was not strongly supported by the data provided in the paper. Three factors influencing how submerged macrophytes respond to high ammonium, not addressed by Yu et al.'s paper, are toxicity of ammonium/ammonia (NH4(+)/NH3), the precise extent of shading in water and species-specific characteristics of macrophytes. In conclusion, a comprehensive consideration of abiotic and biotic factors that involve in the responses of submerged macrophytes to high N is urged in future studies of the role of high N on the growth of submerged macrophytes.
Article
Loss of submersed macrophytes is a world-wide phenomenon occurring when shallow lakes become eutrophic due to excess nutrient loading. In addition to the well-known effect of phosphorus, nitrogen as a trigger of macrophyte decline has received increasing attention. The precise impact of high nitrogen concentrations is debated, and the role of different candidate factors may well change over the season. In this study, we conducted experiments with Vallisneria natans during the growing season (June–September) in 10 ponds subjected to substantial differences in nitrogen loading (five targeted total nitrogen concentrations: control, 2, 10, 20, and 100 mg L− 1) and compared the results with those obtained in our earlier published study from the low-growth season (December–April). Like in the low-growth season, growth of V. natans in summer declined with increasing ammonium (NH4) concentrations and particularly with increasing phytoplankton chlorophyll a (ChlaPhyt). Accordingly, we propose that shading by phytoplankton might be of key importance for macrophyte decline, affecting also periphyton growth as periphyton chlorophyll a (ChlaPeri) decreased with increasing ChlaPhyt. Free amino acid contents (FAA) of plants tended to increase with increasing NH4 concentrations, while the relationships between FAA with growth indices were all weak, suggesting that FAA might be a useful indicator of the physiological stress of the plants but not of macrophyte growth. Taken together, the results from the two seasons indicate that although a combination of high nitrogen concentrations (ammonium) and shading by phytoplankton may cause severe stress on macrophytes, active growth in the growing season enabled them to partly overcome the stress.
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The toxicity of ammonia to fishes has been attributed to the un-ionized ammonia chemical species present in aqueous solution. Because the percent of total ammonia present as un-ionized ammonia (NH3) is so dependent upon pH and temperature, an exact understanding of the aqueous ammonia equilibrium is important for toxicity studies. A critical evaluation of the literature data on the ammonia–water equilibrium system has been carried out. Results of calculations of values of pKa at different temperatures and of percent of NH3 in aqueous ammonia solutions of zero salinity as a function of pH and temperature are presented.
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Water managers often debate whether resuspension of sediment with high organic matter and water content accumulated during eutrophication delays improvement of water clarity after reduction of external nutrient loading. Using data from 15 shallow (mean depth <5 m) eutrophic lakes surveyed during 8-12 yr, we show that the reduction in phytoplankton biomass after external loading reductions of phosphorus or changes in the abundance of plankti-benthivorous fish was accompanied by a proportional or nearly proportional reduction in detritus and inorganic suspended solids. The reduction occurred irrespective of lake size (0.1-40 km(2)), extent of phytoplankton biomass reduction (up to 10-fold), and despite dominance of sediments with high water and organic content. Therefore, we conclude that recovery of shallow lakes after nutrient loading or fish stock reduction is apparently not significantly delayed by resuspension of organic or inorganic matter accumulated in the sediment during eutrophication.
Article
1. While phosphorus (P) is often considered the most important growth limiting factor for plants in lakes, recent studies of shallow lakes indicate that nitrogen (N) may be of greater importance than realized hitherto and that submerged macrophytes may be lost when the N concentration exceeds a certain threshold, as long as the concentration of P is sufficiently high. 2. We studied the effects of different loadings of NH4-N and NO3-N on chlorophyll a and on a macrophyte tolerant of eutrophication, Vallisneria spinulosa (Hydrocharitaceae). In outdoor mesocosms we used water from a pond as control and created four levels of NH4-N and NO3-N (approximately 2.5, 5, 7.5 and 10 mg L−1) by dosing with NH4Cl and NaNO3, respectively. After the experiment, the plants were transferred back to a holding pond to study their recovery. In contrast to previous research, we used a low background concentration of phosphorus (TP 0.024 ± 0.003 mg L−1) so we could judge whether any effects of N were apparent when P is in short supply. 3. Chlorophyll a increased significantly with N dosing for both forms of N, but the increase was highest in the NH4-N dosed mesocosms (maximum 58 μg L−1 versus 42 μg L−1 in the NO3-N mesocosms), probably due to a higher total inorganic N concentration (part of the added ammonia was converted to nitrate during the experiment). 4. Although the number of ramets of V. spinulosa was not affected by the N treatment, the biomass increased up to concentrations of 7.5 mg L−1, while biomass at 10 mg L−1 remained at the control level for both N ions treatments. A similar pattern was apparent for the content of N and soluble sugar of the plant, while there were no differences in the plant P content among treatments. Five months after transplantation back to the pond no difference was found in the number of ramets or in biomass, except that the biomass of plants grown at 10 mg N L−1 during the experiment was greater than that in the control, while the N and P contents of plants were similar to those of the controls. 5. Nitrogen concentration had little influence on the growth of the eutrophication tolerant submerged macrophyte at moderately low concentrations of phosphorus. Moreover, the two N ions showed no toxic effects, suggesting that loss of macrophytes observed in other studies, run at higher phosphorus concentrations, was probably related to enhanced shading by periphyton and/or phytoplankton rather than to any toxic effects of N.
Article
1. Increased ammonium concentrations and decreased light availability in a water column have been reported to adversely affect submersed vegetation in eutrophic waters worldwide. 2. We studied the chronic effects of moderate enrichment (NH4–N: 0.16–0.25 mg L−1) on the growth and carbon and nitrogen metabolism of three macrophytes (Ceratophyllum demersum, Myriophyllum spicatum and Vallisneria natans) under contrasting light availability in a 2-month experiment. 3. The enrichment greatly increased the contents of free amino acids and nitrogen in the shoot / leaf of the macrophytes. This indicates that was the dominant N source for the macrophytes. 4. Soluble carbohydrate contents remained relatively stable in the shoot / leaf of the macrophytes irrespective of the treatments. Under ambient light, the starch contents in the shoot / leaf of C. demersum and M. spicatum increased with enrichment, whereas V. natans did not exhibit any change. The starch contents decreased in C. demersum, increased in M. spicatum and remained unchanged in V. natans after the combined treatment of enrichment and reduced light. 5. The enrichment did not affect the growth of the three macrophytes under the ambient light. However, it did suppress the growth of C. demersum and M. spicatum under the reduced light. The results indicate that a moderate enrichment was not directly toxic to the macrophytes although it might change their viability in eutrophic lakes in terms of the carbon and nitrogen metabolism.
Variations of algal and non-algal turbidity among the Yangtze subtropical shallow lakes
  • H J Wang
  • H Z Wang
  • B Z Pan
  • X M Liang
Wang, H.J., Wang, H.Z., Pan, B.Z., Liang, X.M., 2017. Variations of algal and non-algal turbidity among the Yangtze subtropical shallow lakes. Acta Hydrobiol. Sin. 41, 414-419 (In Chinese with English abstract).