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Abstract Nitrogen limitation has been proven to inhibit Microcystis proliferation, and the significant decline in Microcystis blooms in late summer or autumn has been considered to be related to the nitrogen depletion in water. Sinking loss is another factor that influences the dynamics of cyanobacteria in lakes. However, to date, it is still unclear how the sinking property of Microcystis responds to nitrogen availability. Our results suggest that nitrogen limitation would directly influence sinking property of Microcystis , through a significant increase in the specific density of cells. In the short term, carbohydrate accumulation was mainly responsible for the high specific density, showing a high correlation among the NO 3 −-N concentration, specific density and carbohydrate content. Furthermore, carbohydrates could rapidly accumulate after one light/dark cycle, which was mainly due to the reduction in carbohydrate consumption in the darkness under nitrogen limitation. Under nitrogen-light coupling conditions, the specific density ranged from 1.060 to 1.068, except for the treatment with high-nitrogen plus low-light, which showed the value of 1.032. More importantly, when coupled with low nitrogen, the low light did not decrease the carbohydrate content and the specific density, which implied that the sinking cells could not migrate back to the surface. Accordingly, a hypothesis was proposed that the carbohydrate accumulation induced by low nitrogen availability caused an increase in specific density, which invalidates the buoyancy regulation, and cells sink continually out of the water column. This study explores a new understanding on the disappearance mechanisms of Microcystis blooms in the late summer and fall. Graphical abstract Image 1001798 Highlights • Nitrogen limitation increased the specific density of Microcystis. • The higher specific density was attributed to the accumulation of carbohydrates. • Rapid carbohydrate accumulation was due to its lower consumption rate in darkness. • Nitrogen limitation negatively affected the buoyancy regulation. • This study complements the disappearance mechanisms of Microcystis blooms. [ABSTRACT FROM AUTHOR]

Denitrification is the most important process for nitrogen removal in eutrophic lakes and was mostly investigated in lake sediment. Denitrification could also be mediated by cyanobacterial aggregates, yet how this process impacts nitrogen (N) availability and the associated autotroph-heterotroph relationships within cyanobacterial aggregates has not been investigated. In this study, incubation experiments with nitrate amendment were conducted with Microcystis aggregates (MAs). Measurement of nitrogen contents, 16S rRNA-based microbial community profiling and metatranscriptomic sequencing were used to jointly assess nitrogen turnover dynamics, as well as changes in microbial composition and gene expression. Strong denitrification potential was revealed, and maximal N removal was achieved within two days, after which the communities entered a state of severe N limitation. Changes of active microbial communities were further promoted both with regard to taxonomic composition and transcriptive activities. Expression of transportation-related genes confirmed competition for N sources by Microcystis and phycospheric communities. Strong stress response to reactive oxygen species by Microcystis was revealed. Notably, interspecific relationships among Microcystis and phycospheric communities exhibited a shift toward antagonistic interactions, particularly evidenced by overall increased expression of genes related to cell lysis and utilization of cellular materials. Patterns of fatty acid and starch metabolism also suggested changes in carbon metabolism and cross-feeding patterns within MAs. Taken together, this study demonstrated substantial denitrification potential of MAs, which, importantly, further induced changes in both metabolic activities and autotroph-heterotroph interactions. These findings also highlight the key role of nutrient condition in shaping autotroph-heterotroph relationships. [Display omitted] • Microcystis aggregates have strong denitrification potential. • Denitrification shifted composition and gene expression of active communities. • Prolonged incubation promoted antagonistic autotroph-heterotroph relationships. • Changes in carbon metabolism and cross-feeding patterns within Microcystis aggregates also occurred. [ABSTRACT FROM AUTHOR]

Recent studies indicated that the algal decomposition produces particulate and dissolved organic carbon (DOC), and can enhance denitrification in eutrophic lakes. However, the effects of the living cyanobacteria on nitrogen cycling in eutrophic lakes were still an unknown question. This study explores a new underlying mechanism of nitrate removal which is driven by living Microcystis . The results suggested that living Microcystis significantly enhanced the nitrate removal at sediment-water interface, with a nitrate removal rate of 0.54 d −1 , which was 2.57 times higher than the nitrate removal rate in the treatment without the addition of Microcystis . Measurements of Chl a and Fv/Fm confirmed that Microcystis was tolerant to the dark/anoxic condition, and the recovery experiments suggested that Microcystis could survive under such stress conditions for at least seven days. Meanwhile, DOC secreted by living Microcystis reached to 4.55 mg C mg −1 Chl a. These secretions were biodegradable hydrophilic and contained carbohydrates and proteins. Our study indicated that during blooms, sinking Microcystis cells could directly provide DOC as carbon source, then consequently enhanced the denitrification at sediment-water interface, and the interactive relationship between living cyanobacteria and permanent nitrate removal should be taken into account while studying nitrogen cycling in aquatic ecosystem. [ABSTRACT FROM AUTHOR]

Harmful cyanobacterial blooms (CyanoHABs) represent a serious threat to aquatic ecosystems. A beneficial use for these harmful microorganisms would be a promising resolution of this urgent issue. This study applied a simple method, nitrogen limitation, to cultivate cyanobacteria aimed at producing cyanobacterial carbon for denitrification. Under nitrogen-limited conditions, the common cyanobacterium, Microcystis, efficiently used nitrate, and had a higher intracellular C/N ratio. More importantly, organic carbons easily leached from its dry powder; these leachates were biodegradable and contained a larger amount of dissolved organic carbon (DOC) and carbohydrates, but a smaller amount of dissolved total nitrogen (DTN) and proteins. When applied to an anoxic system with a sediment-water interface, a significant increase of the specific NO X − -N removal rate was observed that was 14.2 times greater than that of the control. This study first suggests that nitrogen-limited cultivation is an efficient way to induce organic and carbohydrate accumulation in cyanobacteria, as well as a high C/N ratio, and that these cyanobacteria can act as a promising carbon source for denitrification. The results indicate that application as a carbon source is not only a new way to utilize cyanobacteria, but it also contributes to nitrogen removal in aquatic ecosystems, further limiting the proliferation of CyanoHABs. [ABSTRACT FROM AUTHOR]

In-situ light-availability control is commonly used to suppress Microcystis blooms in nutrient-rich water resources. It has been suggested that the reduction of column cyanobacterial biomass could mostly be attributed to the inhibition of photosynthesis. However, sinking loss may be another factor influencing the column cyanobacterial biomass. To further investigate the mechanism of this reduction, a mixing-static water column experimental apparatus was designed to simulate the reduction of Microcystis biomass under light-availability control. Under light-shading plus mixing, the reduction of Microcystis in the water column was attributed to both intrinsic biomass loss and sinking loss. Comparatively, under light-shading without mixing, the Microcystis accumulated in surface water, maintaining a continuously increase of intrinsic biomass. Meanwhile, the sinking loss increased as the water column became static, even exceeding the increase of intrinsic biomass, suggesting that sinking loss was the main mechanism for the reduction under light-shading. Further investigation indicated that both intrinsic growth rate and sinking loss rate varied in response to available light. Accordingly, a hypothesis is represented that the loss of column biomass and the shift in dominant species under light-availability control are primarily attributed to the combined effects of intrinsic biomass change and sinking loss, which both respond to available light. [ABSTRACT FROM AUTHOR]

• Microcystis blooms play an important role in N reduction with different mechanisms. • N isotope test indicated nitrate was reduced mainly by denitrification in the bottom. • Microcystis is able to reduce N from the overlying water through denitrification. • 4 Field study also supported the enhancing effect of Microcystis on denitrification. • This study provided insights into the feedback of Microcystis bloom on N cycle. During the bloom seasons, the dissolved inorganic nitrogen declines, which results in the occurrence of nitrogen limitation. It is unclear where the nitrogen goes. Our enclosure experiments and batch tests suggested that Microcystis blooms could significantly reduce the nitrogen in water bodies and the key mechanisms for the nitrogen reduction in different layers were different. The assimilation was the main pathway for nitrogen reduction in the surface layer, while denitrification played an important role both at the sediment-water interface and in the overlying water. Stable nitrogen isotope experiments showed that the nitrate reduction efficiency at sediment-water interface was enhanced by Microcystis , reaching to 76.5∼84.7 %. Dissimilation accounted for 63.8∼67.3 % of the nitrate reduction, and the denitrification rate was 7.4∼8.5 times of DNRA rate. In the water column, the Microcystis bloom facilitated the formation of dark/anoxic condition, which favored the denitrification. The Microcystis aggregates collected from the field showed a great potential in removing nitrogen, and the TN in the overly water was reduced by 3.76∼6.03 mg L−1 within two days. This study provided field evidences and deeper insights into the relationship between Microcystis blooms and nitrogen reduction in the whole water column and gave more details about the enhancing effects of Microcystis on nitrogen reduction. [ABSTRACT FROM AUTHOR]

• Low nitrogen availability promotes the downward transport of Microcystis biomass. • The increased downward transport of biomass is driven by physiological processes. • Carbohydrate accumulation is mainly due to a downregulation of glycolysis. • The increased downward transport of biomass accelerates the dissipation of bloom. Nitrogen availability is one of the key factors affecting the dynamics of non-diazotrophic cyanobacterial blooms in eutrophic lakes. While previous studies mainly focused on the promoting effect of nitrogen on the growth of cyanobacteria, this study aimed to investigate the role of nitrogen availability in the downward transport of biomass and its effects on the dynamics of Microcystis blooms. We performed field enclosure experiments which demonstrated that nitrogen availability negatively affects the downward transport of biomass. With a nitrogen loading of 0.02 g N m−2 d−1, the Microcystis biomass in the water column decreased by 56.2% over a 4-day period. During the same period of time, the average sinking ratio was 0.23 d−1; moreover, the termination of biomass growth was detected. At the notably higher nitrogen loading of 0.5 g N m−2d−1, the downward transport of biomass could still compensate for the biomass growth, although the average sinking ratio was lower at 0.16 d−1. Additional laboratory culture experiments demonstrated that the increase in the downward transport of Microcystis occurred in parallel to an increase in the carbohydrate content and a decrease in gas vesicle content. Further proteomic analysis indicated that the carbohydrate accumulation induced by nitrogen deficiency was a result of the slowing down of catabolic consumption, especially the downregulation of glycolysis. Thus, our study suggests that increased intracellular carbohydrate accumulation at low nitrogen availability causes a higher sinking ratio of Microcystis , indicating that nitrogen limits the duration of Microcystis blooms; thus, decreased nitrogen availability may lead to increased sinking of biomass out of the water column, accelerating the dissipation of Microcystis blooms. [ABSTRACT FROM AUTHOR]

• The salt tolerance threshold of Microcystis showed temporal variation in nature. • Upstream nitrogen availability determines the salt tolerance of Microcystis. • Salt tolerance depends on carbohydrate content and nitrogen availability. • Microcystin content and release depend on the background nitrogen availability. • Natural Microcystis colonies pose a larger risk to brackish water in summer. The occurrence of large Microcystis biomass in brackish waters is primarily caused by its downward transportation from the upstream freshwater lakes and reservoirs through rivers rather than due to in situ bloom formation. Factors that determine the survival of freshwater cyanobacteria in brackish waters have not been well investigated. Here, we studied the spatiotemporal variability of inorganic nitrogen in an upstream lake and conducted laboratory and in-situ experiments to assess the role of nitrogen availability on the salt tolerance of Microcystis and the release of microcystins. A series of field experiments were carried out during bloom seasons to evaluate the salt tolerance of natural Microcystis colonies. The salt tolerance threshold varied from 7 to 17 and showed a positive relationship with intracellular carbohydrate content and a negative relationship with nitrogen availability in water. In August when upstream nitrogen availability was lower, the Microcystis colonies could maintain their biomass even after a sudden increase in salinity from 4 to 10. Laboratory-cultivated Microcystis that accumulated higher carbohydrate content at lower nitrogen availability showed better cell survival at higher salinity. The sharp release of microcystins into the surrounding water occurred when salinity exceeded the salt tolerance threshold of the Microcystis. Thus, Microcystis with higher salt tolerance can accumulate more toxins in cells. The obtained results suggest that the cell survival and toxin concentration in brackish waters depend on the physiological properties of Microcystis formed in the upstream waters. Thus, the life history of Microcystis in upstream waters could have a significant impact on its salt tolerance in downstream brackish waters, where the ecological risk of the salt-tolerant Microcystis requires special and careful management in summer at low nitrogen availability. [Display omitted] [ABSTRACT FROM AUTHOR]