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1. Variable Stoichiometry Effects on Glacial/Interglacial Ocean Model Biogeochemical Cycles and Carbon Storage

   Fillman, Nathaniel (June 2023, Scholar's Archive at OSU)

   Realistic model representation of ocean phytoplankton is important for simulating nutrient cycles and the biological carbon pump, which affects atmospheric carbon dioxide (pCO2) concentrations and, thus, climate. Until recently, most models assumed constant ratios (or stoichiometry) of phosphorous (P), nitrogen (N), silicon (Si), and carbon (C) in phytoplankton, despite observations indicating systematic variations. Here, we investigate the effects of variable stoichiometry on simulated nutrient distributions, plankton community compositions, and the C cycle in the preindustrial (PI) and glacial oceans. Using a biogeochemical model, a linearly increasing P:N relation to increasing PO4 is implemented for ordinary phytoplankton (PO), and a nonlinearly decreasing Si:N relation to increasing Fe is applied to diatoms (PDiat). C:N remains fixed. Variable P:N affects modeled community composition through enhanced PO4 availability, which increases N-fixers in the oligotrophic ocean, consistent with previous research. This increases the NO3 fertilization of PO, the NO3 inventory, and the total plankton biomass. The accuracy of modeled surface nutrients is relatively unchanged. Conversely, variable Si:N shifts south the Southern Ocean’s meridional surface silicate gradient, which aligns better with observations, but depresses PDiat growth globally. In Last Glacial Maximum simulations, PO respond to more oligotrophic conditions by increasing their C:P. This strengthens the biologically mediated C storage such that dissolved organic (inorganic) C inventories increase by 34-40 (38-50) Pg C and 0.7-1.2 Pg yr-1 more particulate C is exported into the interior ocean. Thus, an additional 13-14 ppm of pCO2 difference from PI levels results, improving model agreement with glacial observations. 

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2. Low rates of bacterivory enhances phototrophy and competitive advantage for mixoplankton growing in oligotrophic waters

   https://doi.org/10.1038/s41598-023-33962-x  

   Mitra, Aditee; Flynn, Kevin J. (December 2023, Scientific Reports)

   Abstract With climate change, oceans are becoming increasingly nutrient limited, favouring growth of prokaryotic picoplankton at the expense of the larger protist plankton whose growth support higher trophic levels. Constitutive mixoplankton (CM), microalgal plankton with innate phototrophic capability coupled with phagotrophy, graze on these picoplankton, indirectly exploiting the excellent resource acquisition abilities of the prokaryotes. However, feeding rates can be very low (e.g., a few bacteria d −1 ). For the first time, the significance of such low consumption rates has been quantified. We find that while prokaryote-carbon (C) supply to CM grown at non-limiting light was so low that it may appear insignificant (< 10%), contributions of nitrogen (N) and phosphorus (P) from ingestions of 1–12 prokaryotes d −1 were significant. Under limiting light, contributions of ingested C increased, also raising the contributions of N and P. The order of nutritional importance for CM growth from predation was P > N > C. Further, provision of N through internal recycling of ingested prey-N stimulates C-fixation through photosynthesis. Importantly, coupled photo-phago-mixoplanktonic activity improved CM resource affinities for both inorganic and prey-bound nutrients, enhancing the nutritional status and competitiveness of mixoplankton. With warming oceans, with increased prokaryote abundance, we expect CM to exhibit more phagotrophy. 

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3. Resource limitation of autotrophs and heterotrophs in boreal forest headwater streams

   https://doi.org/10.1086/722256  

   Weaver, Sophie A.; Jones, Jeremy B. (December 2022, Freshwater Science)

   Autotrophic and heterotrophic microbes in stream biofilms dominate biogeochemical cycling and rely on nutrient and energy resources for growth and productivity. In the boreal forest, variation in these resources can originate from permafrost distribution and controls competition for nutrients between stream autotrophs and heterotrophs. We investigated which resources control nutrient uptake and metabolism in headwater stream biofilms of subarctic Alaska, USA, and how resource availability affects competition for inorganic nutrients. We hypothesized that the competitive outcome between autotrophs and heterotrophs for inorganic nutrients would be dependent on availability of organic C, or inorganic nutrients (N and P). To test our hypotheses, we measured resource limitation at the patch and reach scales along a permafrost gradient in interior Alaska. At the patch scale, nutrient diffusing substrata revealed that, secondary to light, N and P were colimiting to autotrophic growth, whereas C was primarily limiting to heterotrophic respiration. In the presence of labile C, heterotrophs exhibited a larger response to nutrient enrichment and outcompeted autotrophs for inorganic nutrients. At the reach scale, light availability had the largest influence on nutrient uptake, but inorganic nutrients were also important. The positive response to increased nutrient and C availability at the patch scale suggests that the predicted increase in exports into fluvial networks with permafrost degradation will alter biofilm structure and function. Ultimately, biofilm communities will shift to more heterotroph-dominated patches if heterotrophs outcompete autotrophs for inorganic nutrients. As permafrost thaws and nutrients and organic C mobilize into streams, nutrient uptake dynamics and competition within biofilms will be altered, affecting nutrient use and export. 

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4. Multiple biotic interactions establish phytoplankton community structure across environmental gradients

   https://doi.org/10.1002/lno.12555  

   Dutkiewicz, Stephanie; Follett, Christopher L; Follows, Michael J; Henderikx‐Freitas, Fernanda; Ribalet, Francois; Gradoville, Mary R; Coesel, Sacha N; Farnelid, Hanna; Finkel, Zoe V; Irwin, Andrew J; et al (May 2024, Limnology and Oceanography)

   Abstract The combination of taxa and size classes of phytoplankton that coexist at any location affects the structure of the marine food web and the magnitude of carbon fluxes to the deep ocean. But what controls the patterns of this community structure across environmental gradients remains unclear. Here, we focus on the North East Pacific Transition Zone, a ~ 10° region of latitude straddling warm, nutrient‐poor subtropical and cold, nutrient‐rich subpolar gyres. Data from three cruises to the region revealed intricate patterns of phytoplankton community structure: poleward increases in the number of cell size classes; increasing biomass of picoeukaryotes and diatoms; decreases in diazotrophs andProchlorococcus; and both increases and decreases inSynechococcus. These patterns can only be partially explained by existing theories. Using data, theory, and numerical simulations, we show that the patterns of plankton distributions across the transition zone are the result of gradients in nutrient supply rates, which control a range of complex biotic interactions. We examine how interactions such as size‐specific grazing, multiple trophic strategies, shared grazing between several phytoplankton size classes and heterotrophic bacteria, and competition for multiple resources can individually explain aspects of the observed community structure. However, it is the combination of all these interactions together that is needed to explain the bulk compositional patterns in phytoplankton across the North East Pacific Transition Zone. The synthesis of multiple mechanisms is essential for us to begin to understand the shaping of community structure over large environmental gradients. 

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5. Mixotrophy in nanoflagellates across environmental gradients in the ocean

   https://doi.org/10.1073/pnas.1814860116  

   Edwards, Kyle F. (February 2019, Proceedings of the National Academy of Sciences)

   Mixotrophy, the combination of autotrophic and heterotrophic nutrition, is a common trophic strategy among unicellular eukaryotes in the ocean. There are a number of hypotheses about the conditions that select for mixotrophy, and field studies have documented the prevalence of mixotrophy in a range of environments. However, there is currently little evidence for how mixotrophy varies across environmental gradients, and whether empirical patterns support theoretical predictions. Here I synthesize experiments that have quantified the abundance of phototrophic, mixotrophic, and heterotrophic nanoflagellates, to ask whether there are broad patterns in the prevalence of mixotrophy (relative to pure autotrophy and heterotrophy), and to ask whether observed patterns are consistent with a trait-based model of trophic strategies. The data suggest that mixotrophs increase in abundance at lower latitudes, while autotrophs and heterotrophs do not, and that this may be driven by increased light availability. Both mixotrophs and autotrophs increase greatly in productive coastal environments, while heterotrophs increase only slightly. These patterns are consistent with a model of resource competition in which nutrients and carbon can both limit growth and mixotrophs experience a trade-off in allocating biomass to phagotrophy vs. autotrophic functions. Importantly, mixotrophy is selected for under a range of conditions even when mixotrophs experience a penalty for using a generalist trophic strategy, due to the synergy between photosynthetically derived carbon and prey-derived nutrients. For this reason mixotrophy is favored relative to specialist strategies by increased irradiance, while at the same time increased nutrient supply increases the competitive ability of mixotrophs against heterotrophs. 

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