Nitrogen's Role in Toxic Algal Bloom Development in Aquatic Ecosystems

References

[1] Anderson, D.M., Alpermann, T.J., Cembella, A.D., Collos, Y., Masseret, E., and Montresor, M. (2012). The globally distributed genus Alexandrium: multifaceted roles in marine ecosystems and impacts on human health. Harmful Algae, 14, 10-35.

[2] Anderson, D.M., Fensin, E., Gobler, C.J., Hoeglund, A.E., Hubbard, K.A., Kulis, D.M., Landsberg, J.H., Lefebvre, K.A., Provoost, P., Richlen, M.L., Smith, J.L., Solow, A.R., and Trainer, V.L. (2021). Marine harmful algal blooms (HABs) in the United States: history, current status and future trends. Harmful Algae, 102, 101975.

[3] Beusen, A.H.W., Doelman, J.C., Van Beek, L.P.H., Van Puijenbroek, P.J.T.M., Mogollón, J.M., Van Grinsven, H.J.M., Stehfest, E., Van Vuuren, D.P., and Bouwman, A.F. (2022). Exploring river nitrogen and phosphorus loading and export to global coastal waters in the Shared Socio-economic pathways. Global Environmental Change, 72, 102426.

[4] Beversdorf, L.J., Miller, T.R., and McMahon, K.D. (2013). The role of nitrogen fixation in cyanobacterial bloom toxicity in a temperate, eutrophic lake. PLoS ONE, 8, e56103.

[5] Bonilla, S., Aubriot, L., Soares, M.C.S., González-Piana, M., Fabre, A., Huszar, V.L., Lürling, M., Antoniades, D., Padisák, J., and Kruk, C. (2012). What drives the distribution of the bloom-forming cyanobacteria Planktothrix agardhii and Cylindrospermopsis raciborskii? FEMS Microbiology Ecology, 79, 594-607.

[6] Karem M Brandenburg 1, Sylke Wohlrab2 3, Uwe John 2 3., Anke Kremp 4, Jacqueline Jerney 4, Bernd Krock2, and Van de Waal1. (2018). Intraspecific trait variation and trade-offs within and across populations of a toxic dinoflagellate. Ecology Letters, 21, 1561-1571.

[7] Bronk, D.A., See, J.H., Bradley, P., and Killberg, L. (2007). DON as a source of bioavailable nitrogen for phytoplankton. Biogeosciences, 4, 283-296.

[8] Carey, R.O. and Migliaccio, K.W. (2009). Contribution of wastewater treatment plant effluents to nutrient dynamics in aquatic systems: a review. Environmental Management, 44, 205-217.

[9] Carpenter, S.R., Caraco, N.F., Correll, D.L., Howarth, R.W., Sharpley, A.N., and Smith, V.H. (1998). Nonpoint pollution of surface waters with phosphorus and nitrogen. Ecological Applications, 8, 559-568.

[10] Collos, Y., Gagne, C., Laabir, M., Vaquer, A., Cecchi, P., and Souchu, P. (2004). Nitrogenous nutrition of Alexandrium catenella (Dinophyceae) in cultures and in Thau lagoon, southern France. Journal of Phycology, 40, 96-103.

[11] Conley, D.J., Paerl, H.W., Howarth, R.W., Boesch, D.F., Seitzinger, S.P., Havens, K.E., Lancelot, C., and Likens, G.E. (2009). Controlling eutrophication: nitrogen and phosphorus. Science, 323, 1014-1015.

[12] Costa, Diogo & Sutter, Caleb & Shepherd, Anita & Jarvie, Helen & Wilson, Henry & Elliott, Jane & Liu, Jian & Macrae, Merrin. (2022). Impact of climate change on catchment nutrient dynamics: insights from around the world. Environmental Reviews. 31. 10.1139/er-2021-0109.

[13] Crumpton, W.G., Stenback, G.A., Fisher, S.W., Stenback, J.Z., and Green, D.I. (2006). Water quality performance of wetlands receiving nonpoint-source nitrogen loads: nitrate and total n itrogen removal efficiency and controlling factors. Journal of Environmental Quality, 45, 1-13.

[14] Dai, M., Luo, Y., Achterberg, E.P., Browning, T.J., Cai, Y., Cao, Z., Chai, F., Chen, B., Church, M.J., Ci, D., Du, C., et al. (2023). Upper ocean biogeochemistry of the oligotrophic North Pacific Subtropical Gyre: from nutrient sources to carbon export. Reviews of Geophysics, 59, e2020RG000720.

[15] Dolman, A.M., Rücker, J., Pick, F.R., Fastner, J., Rohrlack, T., Mischke, U., and Wiedner, C. (2012). Cyanobacteria and cyanotoxins: the influence of nitrogen versus phosphorus. PLoS ONE, 7, e38757

[16] Dorgham, Mohamed. (2014). Effects of Eutrophication. 10.1007/978-94-007-7814-63.

[17] Downing, J.A., Watson, S.B., and McCauley, E. (2001). Predicting cyanobacteria dominance in lakes.

[18] Elser, J.J., Bracken, M.E.S., Cleland, E.E., Gruner, D.S., Harpole, W.S., Hillebrand, H., Ngai, J.T., Seabloom, E.W., Shurin, J.B., and Smith, J.E. (2007). Global analysis of nitrogen and phosphorus limitation of primary producers in freshwater, marine and terrestrial ecosystems. Ecology Letters, 10, 1135-1142.

[19] Flavia Dory, Veronica Nava, Morena Spreafico, Valentina Orlandi, Valentina Soler, Barbara Leoni. Interaction between temperature and nutrients: How does the phytoplankton community cope with the climate change. Science of the total environment, 906(2024) 167566

[20] Finlay, K., Patoine, A., Donald, D.B., Bogard, M.J., and Leavitt, P.R. (2010). Experimental evidence that pollution with urea can degrade water quality in phosphorus-rich lakes of the Northern Great Plains. Limnology and Oceanography, 55, 1213-1230.

[21] Flores, E. and Herrero, A. (2005). Nitrogen assimilation and nitrogen control in cyanobacteria.

[22] Flynn, K.J., Stoecker, D.K., Mitra, A., Raven, J.A., Glibert, P.M., Hansen, P.J., Granéli, E., and Burkholder, J.M. (2013). Misuse of the phytoplankton-zooplankton dichotomy: the need to assign organisms as mixotrophs within plankton functional types. Journal of Plankton Research, 35, 3-11.

[23] Flynn, K.J., Mitra, A., Anestis, K., Anschütz, A.A., Calbet, A., Ferreira, G.D., Gypens, N., Hansen, P.J., John, U., Martin, J.L., Mansour, J.S., et al. (2022). Mixotrophic protists and a new paradigm for marine ecology: where does plankton research go now? Journal of Plankton Research, 41, 375-391.

[24] Fu, F.X., Warner, M.E., Zhang, Y., Feng, Y., and Hutchins, D.A. (2008). Effects of increased temperature and CO₂ on photosynthesis, growth, and elemental ratios in marine Synechococcus and Prochlorococcus (Cyanobacteria). Journal of Phycology, 43, 485-496.

[25] Glibert, P.M. (2016). Margalef revisited: a new phytoplankton mandala incorporating twelve dimensions, including nutritional physiology. Harmful Algae, 55, 25-30.

[26] Glibert, P.M. (2020). Harmful algae at the complex nexus of eutrophication and climate change.

[27] Glibert, P.M., Wilkerson, F.P., Dugdale, R.C., Raven, J.A., Dupont, C.L., Leavitt, P.R., Parker, A.E., Burkholder, J.M., and Kana, T.M. (2016). Pluses and minuses of ammonium and nitrate uptake and assimilation by phytoplankton and implications for productivity and community composition, with emphasis on nitrogen-enriched conditions. Limnology and Oceanography, 61, 165-197.

[28] Glibert, P.M., Heil, C.A., Holligan, P.M., Revilla, M., Hoare, A., Alexander, J., and Mitra, A. (2022). Evidence for dissolved organic nitrogen and phosphorus uptake during a cyanobacterial bloom in Florida Bay. Marine Ecology Progress Series, 280, 73-83.

[29] Gobler, C.J., Burkholder, J.M., Davis, T.W., Harke, M.J., Johengen, T., Stow, C.A., and Van de Waal,

[30] Granéli, E. and Hansen, P.J. (2006). Allelopathy in harmful algae: a mechanism to compete for resources In: Ecology of Harmful Algae (Granéli, E. and Turner, J.T., eds.), pp. 189-201. Springer, Berlin.

[31] Hallegraeff, G.M., Anderson, D.M., Belin, C., Bottein, M.Y.D., Bresnan, E., Chinain, M., Enevoldsen, H., Iwataki, M., Karlson, B., McKenzie, C.H., Sunesen, I., et al. (2021). Perceived global increase in algal blooms is attributable to intensified monitoring and emerging bloom impacts. Communications Earth & Environment, 2, 178.

[32] Justina J. Hampel 1,2*,, Mark J.McCarthy 2, Sanni L Aalto 3, and Silvia E Newel3. (2020). Hurricane disturbance stimulated nitrification and altered ammonia oxidizer community structure in Lake Okeechobee and St. Lucie Estuary (Florida). Frontiers in Microbiology, 10, 2541.

[33] Harke, M.J. and Gobler, C.J. (2013). Global transcriptional responses of the toxic cyanobacterium, Microcystis aeruginosa, to nitrogen stress, phosphorus stress, and growth on organic matter. PLoS ONE, 8, 69834.

[34] Harke, M.J., Steffen, M.M., Gobler, C.J., Otten, T.G., Wilhelm, S.W., Wood, S.A., and Paerl, H.W. (2016). A review of the global ecology, genomics, and biogeography of the toxic cyanobacterium, Microcystis spp. Harmful Algae, 54, 4-20.

[35] K.E Havens 1., T. Fukushima, Xie, P., T. Iwakuma, R.T., James, , N. Takamura, , T. Hanazato, and, T. Yamamoto (2003). Nutrient dynamics and the eutrophication of shallow lakes Kasumigaura (Japan), Donghu (PR China), and Okeechobee (USA). Environmental Pollution, 111, 263-272.

[36] Herbert, E.R., Boon, P., Burgin, A.J., Neubauer, S.C., Franklin, R.B., Ardón, M., Hopfensperger, K.N., Lamers, L.P.M., and Gell, P. (2015). A global perspective on wetland salinization: ecological consequences of a growing threat to freshwater wetlands. Ecosphere, 6, 1-43.

[37] Geoffrey P Horst 1., Orlando Sarnelle 2, Jeffrey D White1, Stephen K. Hamilton 3, Rajreni B Kaul 1., and Juliana D Bressie 4. (2014). Nitrogen availability increases the toxin quota of a harmful cyanobacterium, Microcystis aeruginosa. Water Research, 54, 188-198.

[38] Howarth, R.W. and Marino, R. (2006). Nitrogen as the limiting nutrient for eutrophication in coastal marine ecosystems: evolving views over three decades. Limnology and Oceanography, 51, 364-376.

[39] Huisman, J., Codd G.A., Paerl, H.W., Ibelings, B.W., Verspagen, J.M.H., and Visser, P.M. (2018). Cyanobacterial blooms. Nature Reviews Microbiology, 16, 471-483.

[40] Huisman, J., Sharples, J., Stroom, J.M., Visser, P.M., Kardinaal, W.E.A., Verspagen, J.M.H., and Sommeijer, B. (2004). Changes in turbulent mixing shift competition for light between phytoplankton species. Ecology, 85, 2960-2970.

[41] Hutchins, D.A., Mulholland, M.R., and Fu, F. (2009). Nutrient cycles and marine microbes in a CO₂-enriched ocean. Oceanography, 22, 128-145.

[42] Jeppesen, E., Søndergaard, M., Jensen, J.P., Havens, K.E., Anneville, O., Carvalho, L., Coveney, M.F., Deneke, R., Dokulil, M.T., Foy, B., Gerdeaux, D., et al. (2005). Lake responses to reduced nutrient loading an analysis of contemporary long-term data from 35 case studies. Freshwater Biology, 50, 1747-1771.

[43] John, E.H. and Flynn, K.J. (2010). Growth dynamics and toxicity of Alexandrium fundyense (Dinophyceae): the effect of changing N:P supply ratios on internal toxin and nutrient levels. European Journal of Phycology, 35, 11-23.

[44] Kemp, W.M., Boynton, W.R., Adolf, J.E., Boesch, D.F., Boicourt, W.C., Brush, G., Cornwell, J.C., Fisher, T.R., Glibert, P.M., Hagy, J.D., Harding, L.W., et al. (2005). Eutrophication of Chesapeake Bay: historical trends and ecological interactions. Marine Ecology Progress Series, 303, 1-29.

[45] Klausmeier, C.A., Litchman, E., Daufresne, T., and Levin, S.A. (2004). Optimal nitrogen-to-phosphorus stoichiometry of phytoplankton. Nature, 429, 171-174.

[46] Kosten, S., Huszar, V.L., Bécares, E., Costa, L.S., van Donk, E., Hansson, L.A., Jeppesen, E., Kruk, C., Lacerot, G., Mazzeo, N., De Meester, L., et al. (2012). Warmer climates boost cyanobacterial dominance in shallow lakes. Global Change Biology, 18, 118-126.

[47] Kudela, R.M., Lane, J.Q., and Cochlan, W.P. (2008). The potential role of anthropogenically derived nitrogen in the growth of harmful algae in California, USA. Harmful Algae, 8, 103-110.

[48] Kimberly ALema 1., Maria Latimier 2., Elisabeth Nézan3, Juliette Fauchot 4., and Michael Le Gac, M. (2017). Inter and intra-specific growth and domoic acid production in relation to nutrient ratios and concentrations in Pseudo-nitzschia: phosphate an important factor. Harmful Algae, 64, 11-19.

[49] Li, M., Glibert, P.M., and Gao, Y. (2023). Spatial and temporal dynamics of dissolved inorganic nitrogen in the Chesapeake Bay: a 30-year analysis. Science of the Total Environment, 858, 159774.

[50] Liu, J., Ulén, B., Bergström, L., and Aronsson, H. (2017). Potential phosphorus release from catch crop shoots and roots after freezing-thawing. Plant and Soil, 371, 543-557.

[51] Lomas, M.W. and Glibert, P.M. (1999). Interactions between NH₄⁺ and NO₃⁻ uptake and assimilation: comparison of diatoms and dinoflagellates at several growth temperatures. Marine Biology, 133, 541-551.

[52] Meals, D.W., Dressing, S.A., and Davenport, T.E. (2010). Lag time in water quality response to best management practices: a review. Journal of Environmental Quality, 39, 85-96.

[53] Meissner, S., Fastner, J., and Dittmann, E. (2013). Microcystin production revisited: conjugate formation makes a major contribution. Environmental Microbiology, 15, 1810-1820.

[54] Michalak, A.M., Anderson, E.J., Beletsky, D., Boland, S., Bosch, N.S., Bridgeman, T.B., Chaffin, J.D., Cho, K., Confesor, R., Daloglu, I., DePinto, J.V., et al. (2013). Record-setting algal bloom in Lake Erie caused by agricultural and meteorological trends consistent with expected future conditions. Proceedings of the National Academy of Sciences, 110, 6448-6452.

[55] Mitsch, W.J., Day, J.W., Gilliam, J.W., Groffman, P.M., Hey, D.L., Randall, G.W., and Wang, N. (2001). Reducing nitrogen loading to the Gulf of Mexico from the Mississippi River Basin: strategies to counter a persistent ecological problem. BioScience, 51, 373-388.

[56] Mosley, L.M. (2015). Drought impacts on the water quality of freshwater systems; review and integration. Earth-Science Reviews, 140, 203-214.

[57] Neilan, B.A., Pearson, L.A., Muenchhoff, J., Moffitt, M.C., and Dittmann, E. (2013). Environmental conditions that influence toxin biosynthesis in cyanobacteria. Environmental Microbiology, 15, 1239-1253.

[58] Neubauer, S.C. (2013). Ecosystem responses of a tidal freshwater marsh experiencing saltwater intrusion and altered hydrology. Estuaries and Coasts, 36, 491-507.

[59] Paerl, H.W. and Huisman, J. (2008). Blooms like it hot. Science, 320, 57-58.

[60] Paerl, H.W. and Otten, T.G. (2013). Harmful cyanobacterial blooms: causes, consequences, and controls. Microbial Ecology, 65, 995-1010.

[61] Paerl, H.W. and Paul, V.J. (2012). Climate change: links to global expansion of harmful cyanobacteria. Water Research, 46, 1349-1363.

[62] Paerl, H.W., Fulton, R.S., Moisander, P.H., and Dyble, J. (2001). Harmful freshwater algal blooms, with an emphasis on cyanobacteria. The Scientific World JOURNAL, 1, 76-113.

[63] Paerl, H.W., Dennis, R.L., and Whitall, D.R. (2006). Atmospheric deposition of nitrogen: implications for nutrient over-enrichment of coastal waters. Estuaries and Coasts, 25, 677-693.

[64] Paerl, H.W., Xu, H., McCarthy, M.J., Zhu, G., Qin, B., Li, Y., and Gardner, W.S. (2011). Controlling harmful cyanobacterial blooms in a hyper-eutrophic lake (Lake Taihu, China): the need for a dual nutrient (N & P) management strategy. Water Research, 45, 1973-1983.

[65] Paerl, H.W., Hall, N.S., and Calandrino, E.S. (2016). Controlling harmful cyanobacterial blooms in a world experiencing anthropogenic and climatic-induced change. Science of the Total Environment, 409, 1739-1745.

[66] Hans.W. Paerl, Gardner, W.S., Havens, K.E., Joyner, A.R., McCarthy, M.J., Newell, S.E., Qin, B., and Scott, J.T. (2020). Mitigating cyanobacterial harmful algal blooms in aquatic ecosystems impacted by climate change and anthropogenic nutrients. Harmful Algae, 54, 213-222.

[67] Elif Pehlivanoglu-Mantas, and David L Sedlak, (2008). Measurement of dissolved organic nitrogen forms in wastewater effluents: concentrations, size distribution and NDMA formation potential. Water Research, 42, 3890-3898.

[68] Juliana S.M. Pimentel and Alesander Giani, . (2014). Microcystin production and regulation under nutrient stress conditions in toxic Microcystis strains. Applied and Environmental Microbiology, 80, 5836-5843.

[69] Pryor, S.C., Howe, J.A., and Kunkel, K.E. (2009). How spatially coherent and statistically robust are temporal changes in extreme precipitation in the contiguous USA? International Journal of Climatology, 29, 31-45.

[70] Rabalais, N.N., Díaz, R.J., Levin, L.A., Turner, R.E., Gilbert, D., and Zhang, J. (2009). Dynamics and distribution of natural and human-caused hypoxia. Biogeosciences, 7, 585-619.

[71] Reichwaldt, E.S. and Ghadouani, A. (2012). Effects of rainfall patterns on toxic cyanobacterial blooms in a changing climate: between simplistic scenarios and complex dynamics. Water Research, 46, 1372-1393.

[72] Reynolds, C.S. (2006). The Ecology of Phytoplankton. Cambridge University Press, Cambridge.

[73] Rigosi, A., Carey, C.C., Ibelings, B.W., and Brookes, J.D. (2014). The interaction between climate warming and eutrophication to promote cyanobacteria is dependent on trophic state and varies among taxa. Limnology and Oceanography, 59, 99-114.

[74] Rohini Mattoo*, Usman B. Mallikarjuna, and Naveenachar Hemachar. Ecosystem and climate change impacts on the nitrogen cycle and biodiversity. Nitrogen, 2025: 6(5), 78

[75] Schindler, D.W., Hecky, R.E., Findlay, D.L., Stainton, M.P., Parker, B.R., Paterson, M.J., Beaty, K.G., Lyng, M., and Kasian, S.E.M. (2008). Eutrophication of lakes cannot be controlled by reducing nitrogen input: results of a 37-year whole-ecosystem experiment. Proceedings of the National Academy of Sciences, 105, 11254-11258.

[76] Schindler, D.W., Carpenter, S.R., Chapra, S.C., Hecky, R.E., and Orihel, D.M. (2016). Reducing phosphorus to curb lake eutrophication is a success. Environmental Science & Technology, 50, 8923-8929.

[77] Scott, J.T. and Grantz, E.M. (2013). N₂ fixation exceeds internal nitrogen loading as a phytoplankton nutrient source in perpetually nitrogen-limited reservoirs. Freshwater Science, 32, 849-861.

[78] Scott, J.T. and Marcarelli, A.M. (2012). Cyanobacteria in freshwater benthic environments. In: Ecology of Cyanobacteria II, pp. 271-289. Springer, Dordrecht.

[79] Seitzinger, S.P., Sanders, R.W., and Styles, R. (2002). Bioavailability of DON from natural and anthropogenic sources to estuarine plankton. Limnology and Oceanography, 47, 353-366.

[80] Seitzinger, S.P., Mayorga, E., Bouwman, A.F., Kroeze, C., Beusen, A.H.W., Billen, G., Van Drecht, G., Dumont, E., Fekete, B.M., Garnier, J., and Harrison, J.A. (2010). Global river nutrient export: a scenario analysis of past and future trends. Global Biogeochemical Cycles, 24, GB0A08.

[81] Sharpley, A.N., Bergström, L., Aronsson, H., Bechmann, M., Bolster, C.H., Börling, K., Djodjic, F., Jarvie, H.P., Schoumans, O.F., Stamm, C., Tonderski, K.S., et al. (2013). Future agriculture with minimized phosphorus losses to waters: research needs and direction. Ambio, 44, 163-179.

[82] Shatwell, T., Thiery, W., and Kirillin, G. (2019). Future projections of temperature and mixing regime of European temperate lakes. Hydrology and Earth System Sciences, 23, 1533-1551.

[83] Smith, V.H. (2003). Eutrophication of freshwater and coastal marine ecosystems a global problem. Environmental Science and Pollution Research, 10, 126-139.

[84] Smith, J.M., Chavez, F.P., and Francis, C.A. (2014). Ammonium uptake by phytoplankton regulates nitrification in the sunlit ocean. PLoS ONE, 9, e108173.

[85] Søndergaard, M., Jensen, J.P., and Jeppesen, E. (2003). Role of sediment and internal loading of phosphorus in shallow lakes. Hydrobiologia, 506, 135-145.

[86] Sommer, U., Adrian, R., De Senerpont Domis, L., Elser, J.J., Gaedke, U., Ibelings, B., Jeppesen, E., Lürling, M., Molinero, J.C., Mooij, W.M., van Donk, E., and Winder, M. (2012). Beyond the Plankton Ecology Group (PEG) Model: Mechanisms driving plankton succession. Annual Review of Ecology, Evolution, and Systematics, 43, 429-448.

[87] Spilling, K., Olli, K., Lehtoranta, J., Kremp, A., Tedesco, L., Tamelander, T., Klais, R., Peltonen, H., and Tamminen, T. (2018). Shifting diatom-dinoflagellate dominance during spring bloom in the Baltic Sea and its potential effects on biogeochemical cycling. Frontiers in Marine Science, 5, 327.

[88] Steffen, M.M., Davis, T.W., McKay, R.M.L., Bullerjahn, G.S., Krausfeldt, L.E., Stough, J.M.A., Neitzey, M.L., Gilbert, N.E., Boyer, G.L., Johengen, T.H., Gossiaux, D.C., et al. (2017). Ecophysiological examination of the Lake Erie Microcystis bloom in 2014: linkages between biology and the water supply shutdown of Toledo, OH. Environmental Science & Technology, 51, 6745-6755.

[89] Steffen, M.M., Belisle, B.S., Watson, S.B., Boyer, G.L., and Wilhelm, S.W. (2014). Status, causes and controls of cyanobacterial blooms in Lake Erie. Journal of Great Lakes Research, 40, 215-225.

[90] Sterner, R.W. (2008). On the phosphorus limitation paradigm for lakes. International Review of Hydrobiology, 93, 433-445.

[91] Stoecker, D.K. and Lavrentyev, P.J. (2018). Mixotrophic plankton in the polar seas: a pan-Arctic review. Frontiers in Marine Science, 5, 292.

[92] Stow, C.A., Cha, Y., Johnson, L.T., Confesor, R., and Richards, R.P. (2015). Long-term and seasonal trend decomposition of Maumee River nutrient inputs to western Lake Erie. Environmental Science & Technology, 49, 3263-3986.

[93] Terseleer, N., Bruggeman, J., Lancelot, C., and Gypens, N. (2014). Trait-based representation of diatom functional diversity in a plankton functional type model of the eutrophied southern North Sea. Limnology and Oceanography, 59, 1958-1972.

[94] Tonitto, C., David, M.B., and Drinkwater, L.E. (2006). Replacing bare fallows with cover crops in fertilizer-intensive cropping systems: a meta-analysis of crop yield and N dynamics. Agriculture, Ecosystems & Environment, 112, 58-72.

[95] Trainer, V.L., Bates, S.S., Lundholm, N., Thessen, A.E., Cochlan, W.P., Adams, N.G., and Trick, C.G. (2012). Pseudo-nitzschia physiological ecology, phylogeny, toxicity, monitoring and impacts on ecosystem health. Harmful Algae, 14, 271-300.

[96] Trainer, V.L., Moore, S.K., Hallegraeff, G., Kudela, R.M., Clement, A., Mardones, J.I., and Cochlan, W.P. (2020). Pelagic harmful algal blooms and climate change: lessons from nature's experiments with extremes. Harmful Algae, 91, 101591.

[97] Trolle, D., Hamilton, D.P., Pilditch, C.A., Duggan, I.C., and Jeppesen, E. (2010). Predicting the effects of climate change on trophic status of three morphologically varying lakes: implications for lake restoration and management. Environmental Modelling & Software, 26, 354-370.

[98] Tully, K., Gedan, K., Epanchin-Niell, R., Strong, A., Bernhardt, E.S., BenDor, T., Mitchell, M., Kominoski, J., Jordan, T.E., Neubauer, S.C., and Weston, N.B. (2019). The invisible flood: the chemistry, ecology, and social implications of coastal saltwater intrusion. BioScience, 69, 368-378.

[99] Van Meter, K.J., Basu, N.B., and Van Cappellen, P. (2016). Two centuries of nitrogen dynamics: legacy sources and sinks in the Mississippi and Susquehanna River Basins. Global Biogeochemical Cycles, 31, 2-23.

[100] Van Meter, K.J., Van Cappellen, P., and Basu, N.B. (2018). Legacy nitrogen may prevent achievement of water quality goals in the Gulf of Mexico. Science, 360, 427-430.

[101] Veraart, A.J., de Klein, J.J.M., and Scheffer, M. (2011). Warming can boost denitrification disproportionately due to altered oxygen dynamics. PLoS ONE, 6, e18508.

[102] Viancelli A, Michelon W. Climate Change and Nitrogen Dynamics: Challenges and Strategies for a sustainable Future. Nitrogen. 2024:5(3) : 688-701.

[103] Visser, P.M., Verspagen, J.M.H., Sandrini, G., Stal, L.J., Matthijs, H.C.P., Davis, T.W., Paerl, H.W., and Huisman, J. (2016). How rising CO₂ and global warming may stimulate harmful cyanobacterial blooms. Harmful Algae, 54, 145-159.

[104] Wells, M.L., Karlson, B., Wulff, A., Kudela, R., Trick, C., Asnaghi, V., Berdalet, E., Cochlan, W., Davidson, K., De Rijcke, M., Dutkiewicz, S., et al. (2020). Future HAB science: directions and challenges in a changing climate. Harmful Algae, 91, 101632.

[105] Wells ML, Trainer VL, Smayda TJ, Karlson BS, Trick CG, Kudela RM, Ishikwa A, Bernard S, Wulff A, Anderson DM, Cochlan WP. (2015). Harmful algal blooms and climate change; Learning from the past and present to forecast the future. harmful Algae. 1(49): 68-93

[106] Withers, P.J.A., Neal, C., Jarvie, H.P., and Doody, D.G. (2014). Agriculture and eutrophication: where do we go from here? Sustainability, 6, 5853-5875.

[107] Woolway, R.I. and Merchant, C.J. (2019). Worldwide alteration of lake mixing regimes in response to climate change. Nature Geoscience, 12, 271-276.

@article{1717270,
      author = {Donald Ugbong Ugbe, Victoria Chioma Ayozie-Samuel, Ernest, Chidozie Iheanyi, Frank Chibuzor Okenmuo, Linnet Nyokabi Wahome; Bello Aliyu},
      title = {Nitrogen's Role in Toxic Algal Bloom Development in Aquatic Ecosystems},
      journal = {Iconic Research And Engineering Journals},
      year = {2026},
      volume = {9},
      number = {11},
      pages = {292-310},
      issn = {2456-8880},
      url = {https://www.irejournals.com/formatedpaper/1717270.pdf},
      abstract = {Harmful algal blooms (HABs) constitute one of the most pressing water quality challenges confronting aquatic ecosystems globally, with nitrogen pollution serving as a fundamental driver of bloom development and persistence. This comprehensive review examines the multifaceted relationships between nitrogen availability and toxic algal bloom formation across freshwater and marine environments. Anthropogenic nitrogen inputs from agricultural runoff, wastewater discharge, and atmospheric deposition have fundamentally altered nutrient dynamics in water bodies worldwide, creating conditions that systematically favor bloom-forming species over diverse phytoplankton communities. The chemical form of nitrogen, whether nitrate, ammonium, or organic compounds differentially influences algal community composition, with various taxa exhibiting distinct preferences and competitive advantages under specific nitrogen regimes. Temporal patterns of nitrogen delivery, from pulsed agricultural runoff to chronic wastewater discharge, shape seasonal bloom dynamics and species succession patterns. Interactions between nitrogen and phosphorus create complex co-limitation scenarios that determine which organisms dominate under given environmental conditions, while nitrogen availability directly influences cellular toxin production in cyanobacteria, dinoflagellates, and diatoms through metabolic regulatory pathways. Climate change intensifies these nitrogen-driven processes through warming, altered precipitation patterns, and strengthened water column stratification. Management strategies encompassing agricultural best practices, wetland restoration, and advanced wastewater treatment show promise for bloom mitigation, though implementation faces substantial economic, political, and technical challenges. The slow recovery of eutrophic systems due to legacy sediment nutrients demands sustained commitment to load reduction over decadal timescales.},
      keywords = {Algae Bloom, Eutrophication, Cyanobacteria, climate change.},
      month = {May},
      doi = {https://doi.org/10.64388/IREV9I11-1717270}
  }