Preventing harmful algal blooms (HABs) is needed to protect lakes and downstream ecosystems. Traditionally, reducing phosphorus (P) inputs was the prescribed solution for lakes, based on the assumption that P universally limits HAB formation. Reduction of P inputs has decreased HABs in many lakes, but was not successful in others. Thus, the "P-only" paradigm is overgeneralized. Whole-lake experiments indicate that HABs are often stimulated more by combined P and nitrogen (N) enrichment rather than N or P alone, indicating that the dynamics of both nutrients are important for HAB control. The changing paradigm from P-only to consideration of dual nutrient control is supported by studies indicating that (1) biological N fixation cannot always meet lake ecosystem N needs, and (2) that anthropogenic N and P loading has increased dramatically in recent decades. Sediment P accumulation supports long-term internal loading, while N may escape via denitrification, leading to perpetual N deficits. Hence, controlling both N and P inputs will help control HABs in some lakes and also reduce N export to downstream N-sensitive ecosystems. Managers should consider whether balanced control of N and P will most effectively reduce HABs along the freshwater-marine continuum.

Cyanobacterial harmful algal blooms (CyanoHABs) are enhanced by anthropogenic pressures, including excessive nutrient (nitrogen, N, and phosphorus, P) inputs and a warming climate. Severe eutrophication in aquatic systems is often manifested as non-N-fixing CyanoHABs (e.g., Microcystis spp.), but the biogeochemical relationship between N inputs/dynamics and CyanoHABs needs definition. Community biological ammonium (NH) demand (CBAD) relates N dynamics to total microbial productivity and NH deprivation in aquatic systems. A mechanistic conceptual model was constructed by combining nutrient cycling and CBAD observations from a spectrum of lakes to assess N cycling interactions with CyanoHABs. Model predictions were supported with CBAD data from a Microcystis bloom in Maumee Bay, Lake Erie, during summer 2015. Nitrogen compounds are transformed to reduced, more bioavailable forms (e.g., NH and urea) favored by CyanoHABs. During blooms, algal biomass increases faster than internal NH regeneration rates, causing high CBAD values. High turnover rates from cell death and remineralization of labile organic matter consume oxygen and enhance denitrification. These processes drive eutrophic systems to NH limitation or colimitation under warm, shallow conditions and support the need for dual nutrient (N and P) control.

Western Lake Erie cyanobacterial harmful algal blooms (cyanoHABs) occur every summer as a result of anthropogenic nutrient loading. Although the physiological importance of nitrogen (N) in supporting bloom biomass and toxin production is established, the role of internal N recycling in the water column to support bloom maintenance is not as well understood. Over three field seasons (2015-2017), we collected water from western Lake Erie and employed bottle incubations with 15 N-ammonium (NH þ 4 ) enrichments to determine NH þ 4 regeneration and potential uptake rates in the water column. Potential NH þ 4 uptake rates followed spatial and seasonal patterns, with greatest rates measured nearest the Maumee River inflow and during peak bloom months (August and September). Regeneration followed a similar spatial pattern but was greatest in early summer (June and July) and supported $ 20-60% of potential NH þ 4 demand during the height of the bloom. Basinwide internal NH þ 4 regeneration during the April-October period could supply NH þ 4 at 60-200% of annual external N loading to the western basin. These results help explain how non-N-fixing cyanoHABs in Lake Erie and other large, eutrophic lakes continue producing biomass and N-rich toxins long after spring nutrient loads are exhausted or transported to other areas. Internal N loads are ultimately driven by external N loads; in low precipitation years, external nutrient loads result in smaller blooms, producing less substrate for subsequent internal N loads. Overall, these findings, along with others, confirm that both internal and external N loading must be considered when evaluating cyanoHAB management strategies.

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