Kristian Spilling

• Bansfield, D.; Spilling, Kristian; Mikola, A. & Piiparinen, J. (2024). An investigation into the ability of three fungi and one yeast to grow and capture nutrients in cheese whey. Bioresource Technology Reports. ISSN 2589-014X. 26, p. 1–7. doi: 10.1016/j.biteb.2024.101854.
• Goldenberg, Silvan Urs; Spisla, Carsten; Sanchez Puerto, Nicolas; Taucher, Jan; Spilling, Kristian & Sswat, Michael [Show all 12 contributors for this article] (2024). Diatom-mediated food web functioning under ocean artificial upwelling. Scientific Reports. ISSN 2045-2322. 14(1), p. 1–10. doi: 10.1038/s41598-024-54345-w.
• Spilling, Kristian; Arellano San Martín, Mirian; Granlund, Mira; Schulz, Kai G.; Spisla, Carsten & Vanharanta, Mari [Show all 8 contributors for this article] (2023). Microzooplankton communities and their grazing of phytoplankton under artificial upwelling in the oligotrophic ocean. Frontiers in Marine Science. ISSN 2296-7745. 10. doi: 10.3389/fmars.2023.1286899. Full text in Research Archive Show summary

  Ocean artificial upwelling has been suggested to boost primary production and increase harvestable resources such as fish. Yet, for this ecosystem-based approach to work, an effective energy transfer up the food web is required. Here, we studied the trophic role of microzooplankton under artificial upwelling via biomass and community composition as well as grazing rates on phytoplankton. Using mesocosms in the oligotrophic ocean, we supplied nutrient-rich deep water at varying intensities (low to high) and addition modes (a Singular large pulse or smaller Recurring pulses). Deep-water fertilization created a diatom-dominated bloom that scaled with the amount of inorganic nutrients added, but also Synechococcus-like cells, picoeukaryotes and nanophytoplankton increased in abundance with added nutrients. After 30 days, towards the end of the experiment, coccolithophores bloomed under recurring upwelling of high intensity. Across all upwelling scenarios, the microzooplankton community was dominated by ciliates, dinoflagellates (mixo- and heterotrophic) and radiolarians. Under the highest upwelling intensity, the average grazing rates of Synechococcus-like cells, picoeukaryotes and nanophytoplankton by microzooplankton were 0.35 d-1 ± 0.18 (SD), 0.09 d-1 ± 0.12 (SD), and 0.11 d-1 ± 0.13 (SD), respectively. There was little temporal variation in grazing of nanophytoplankton but grazing of Synechococcus-like cells and picoeukaryotes were more variable. There were positive correlations between abundance of these groups and grazing rates, suggesting a response in the microzooplankton community to prey availability. The average phytoplankton to microzooplankton ratio (biovolume) increased with added deep water, and this increase was highest in the Singular treatment, reaching ~30 (m3 m-3), whereas the phytoplankton to total zooplankton biomass ratio (weight) increased from ~1 under low upwelling to ~6 (g g-1) in the highest upwelling but without a difference between the Singular and the Recurring mode. Several smaller, recurring upwelling events increased the importance of microzooplankton compared with one large pulse of deep water. Our results demonstrate that microzooplankton would be an important component for trophic transfer if artificial upwelling would be carried out at scale in the oligotrophic ocean.

• Bansfield, D.; Spilling, Kristian; Mikola, A. & Piiparinen, J. (2023). Growth of fungi and yeasts in food production waste streams: a feasibility study. BMC Microbiology. ISSN 1471-2180. 23(1). doi: 10.1186/s12866-023-03083-6. Full text in Research Archive
• Vanharanta, Mari & Spilling, Kristian (2023). Uptake of excess phosphate at low inorganic N:P ratio in a coastal sea afflicted with eutrophication. Marine Ecology Progress Series. ISSN 0171-8630. 718, p. 23–37. doi: 10.3354/meps14391. Show summary

  Following the spring bloom in the northern Baltic Sea, nitrogen limits phytoplankton growth and there is typically a residual phosphate concentration (>0.2 µmol l-1) remaining that is often assumed to induce the recurring blooms of nitrogen-fixing cyanobacteria. However, these cyanobacterial blooms typically occur 2-3 mo later during the summer, when the phosphate concentration has been depleted, and it is unclear what organisms take up the excess phosphate. We studied the removal potential of excess phosphate (0.55 µmol l-1) at different temperatures (10, 13, 16°C) and with or without nitrogen addition in an indoor 20 l tank experiment. In addition, we followed the element pools and plankton community composition. As expected, the phosphate uptake rate was up to 3-fold faster in nitrogen-amended than non-amended tanks, but complete drawdown of phosphate also occurred under severe nitrogen limitation. The uptake ratio of dissolved inorganic nitrogen to phosphorus was 4.6, which is substantially lower than the Redfield ratio (16) and indicates excessive phosphate removal potential relative to nitrogen. A large part of the excess phosphate ended up in the particulate pool, which has a higher potential to sink out from the surface. The nitrogen-fixing cyanobacteria Nodularia spumigena grew close to summer bloom concentrations only in the highest experimental temperature. However, the combined biovolume of all 3 major bloom-forming cyanobacteria accounted for only 5.3% of the total autotropic biovolume, and their potential phosphate uptake was calculated to be <3% of the excess phosphate available at the beginning of the study. Therefore, our results demonstrate that the contribution of filamentous cyanobacteria to the removal of excess phosphate is small.

• Spilling, Kristian; Piiparinen, Jonna; Achterberg, Eric P.; Arístegui, Javier; Bach, Lennart T. & Camarena-Gómez, Maria T. [Show all 13 contributors for this article] (2023). Extracellular enzyme activity in the coastal upwelling system off Peru: A mesocosm experiment. Biogeosciences. ISSN 1726-4170. 20(8), p. 1605–1619. doi: 10.5194/bg-20-1605-2023. Full text in Research Archive
• Spilling, Kristian; Heinemann, Malte; Vanharanta, Mari; Baumann, Moritz; Noche Ferreira, Andrea & Suessle, Philipp [Show all 7 contributors for this article] (2023). Respiration rate scales inversely with sinking speed of settling marine aggregates. PLOS ONE. ISSN 1932-6203. 18(3). doi: 10.1371/journal.pone.0282294. Full text in Research Archive Show summary

  Sinking marine aggregates have been studied for a long time to understand their role in carbon sequestration. Traditionally, sinking speed and respiration rates have been treated as independent variables, but two recent papers suggest that there is a connection albeit in contrasting directions. Here we collected recently formed (<2 days old) aggregates from sediment traps mounted underneath mesocosms during two different experiments. The mesocosms were moored off Gran Canaria, Spain (~ 27.9 N; 15.4 E) in a coastal, sub-tropical and oligotrophic ecosystem. We determined the respiration rates of organisms (mainly heterotrophic prokaryotes) attached to aggregates sinking at different velocities. The average respiration rate of fast sinking aggregates (>100 m d-1) was 0.12 d-1 ± 0.08 d-1 (SD). Slower sinking aggregates (<50 m d-1) had on average higher (p <0.001) and more variable respiration rates (average 0.31 d-1 ± 0.16 d-1, SD). There was evidence that slower sinking aggregates had higher porosity than fast sinking aggregates, and we hypothesize that higher porosity increase the settlement area for bacteria and the respiration rate. These findings provide insights into the efficiency of the biological carbon pump and help resolve the apparent discrepancy in the recent studies of the correlation between respiration and sinking speed.

• Spilling, Kristian (2022). Bioflocculation of Euglena gracilis via direct application of fungal filaments: a rapid harvesting method. Journal of Applied Phycology. ISSN 0921-8971. doi: 10.1007/s10811-021-02651-5.
• Spilling, Kristian; Asmala, Eero; Haavisto, Noora; Haraguchi, Lumi; Kraft, Kaisa & Lehto, Anne-Mari [Show all 14 contributors for this article] (2022). Dataset from a mesocosm experiment on brownification in the Baltic Sea. Data in Brief. ISSN 2352-3409. 45(108755). doi: 10.1016/j.dib.2022.108755. Full text in Research Archive Show summary

  Climate change is projected to cause brownification of some coastal seas due to increased runoff of terrestrially derived organic matter. We carried out a mesocosm experiment over 15 days to test the effect of this on the planktonic ecosystem. The experiment was set up in 2.2 m3 plastic bags moored outside the Tvärminne Zoological Station at the SW coast of Finland. We used four treatments, each with three replicates: control (Contr) without any manipulation; addition of a commercially available organic carbon additive called HuminFeed (Hum; 2 mg L−1); addition of inorganic nutrients (Nutr; 5.7 µM NH4 and 0.65µM PO4); and a final treatment of combined Nutr and Hum (Nutr+Hum) additions. Water samples were taken daily, and measured variables included water transparency, organic and inorganic nutrient pools, chlorophyll a (Chla), primary and bacterial production and particle counts by flow cytometry.

• Paul, Allanah Joy; Bach, Lennart Thomas; Arístegui, Javier; Von Der Esch, Elisabeth; Hernández-Hernández, Nauzet & Piiparinen, Jonna [Show all 9 contributors for this article] (2022). Upwelled plankton community modulates surface bloom succession and nutrient availability in a natural plankton assemblage. Biogeosciences. ISSN 1726-4170. 19(24), p. 5911–5926. doi: 10.5194/bg-19-5911-2022. Full text in Research Archive Show summary

  Upwelling of nutrient-rich waters into the sunlit surface layer of the ocean supports high primary productivity in eastern boundary upwelling systems (EBUSs). However, subsurface waters contain not only macronutrients (N, P, Si) but also micronutrients, organic matter and seed microbial communities that may modify the response to macronutrient inputs via upwelling. These additional factors are often neglected when investigating upwelling impacts on surface ocean productivity. Here, we investigated how different components of upwelled water (macronutrients, organic nutrients and seed communities) drive the response of surface plankton communities to upwelling in the Peruvian coastal zone. Results from our short-term (10 d) study show that the most influential drivers in upwelled deep water are (1) the ratio of inorganic nutrients (NOx : PO) and (2) the microbial community present that can seed heterogeneity in phytoplankton succession and modify the stoichiometry of residual inorganic nutrients after phytoplankton blooms. Hence, this study suggests that phytoplankton succession after upwelling is modified by factors other than the physical supply of inorganic nutrients. This would likely affect trophic transfer and overall productivity in these highly fertile marine ecosystems.

• Spilling, Kristian; Asmala, Eero; Haavisto, Noora; Haraguchi, Lumi; Kraft, Kaisa & Lehto, Anne-Mari [Show all 14 contributors for this article] (2022). Brownification affects phytoplankton community composition but not primary productivity in eutrophic coastal waters: A mesocosm experiment in the Baltic Sea. Science of the Total Environment. ISSN 0048-9697. 841, p. 1–9. doi: 10.1016/j.scitotenv.2022.156510. Show summary

  Climate change is projected to cause brownification of some coastal seas due to increased runoff of terrestrially derived organic matter. We carried out a mesocosm experiment (15 d) to test the effect of this on the planktonic ecosystem ex- pecting reduced primary production and shifts in the phytoplankton community composition. The experiment was set up in 2.2 m3 mesocosm bags using four treatments, each with three replicates: control (Contr) without any manipula- tion, organic carbon additive HuminFeed (Hum; 2 mg L−1), inorganic nutrients (Nutr; 5.7 μM NH4 and 0.65 μM PO4), and combined Nutr and Hum (Nutr + Hum) additions. Measured variables included organic and inorganic nutrient pools, chlorophyll a (Chla), primary and bacterial production and particle counts by flow cytometry. The bags with added inorganic nutrients developed a phytoplankton bloom that depleted inorganic N at day 6, followed by a rapid decline in Chla. Brownification did not reduce primary production at the tested concentration. Bacterial production was lowest in the Contr, but similar in the three treatments receiving additions likely due to increased carbon available for heterotrophic bacteria. Picoeukaryotes clearly benefited by brownification after inorganic N depletion, which could be due to more effective nutrient recycling, nutrient affinity, light absorption, or alternatively lower grazing pressure. In conclusion, brownification shifted the phytoplankton community composition towards smaller species with potential effects on carbon fluxes, such as sinking rates and export to the sea floor.

• Camarena-Gómez, María Teresa; Lähteenmäki-Uutela, Anu & Spilling, Kristian (2022). Macroalgae production in Northern Europe: Business and government perspectives on how to regulate a novel blue bioeconomy. Aquaculture. ISSN 0044-8486. 560. doi: 10.1016/j.aquaculture.2022.738434. Full text in Research Archive Show summary

  Macroalgae biomass production, understood as cultivation and harvesting, is a minor industry in Europe at present, but the sector is recognized as having substantial growth potential. Here, we framed the environmental license as a boundary object between business and authorities and investigated the details of macroalgal licensing procedures in seven Northern European countries (Finland, Estonia, Sweden, Germany, Norway, Ice- land, and Scotland). We conducted surveys and interviews with macroalgae companies and licensing authorities to understand the challenges faced by both sides. Generally, macroalgae production in Northern European countries is regulated by environmental and water laws and is not included in maritime spatial plans. Private actors need to apply for an environmental, water and/ or fishing permit to start operations in this sector, often with several authorities involved. The companies expressed their dissatisfaction with non-specific laws and burdensome licensing procedures that may delay or even prevent the start-up. The authorities highlighted the lack of scientific environmental risk assessments of macroalgae production and the need to resolve possible conflicts with other marine sectors. Companies need the license to access markets while authorities view the license as a tool to enforce environmental legislation. As a boundary object, the license is the result of correspondence and cooperation between companies and authorities. A one-step licensing procedure for macroalgae production should be applied to encourage this business and to facilitate the compliance of legislation. In addition, macroalgae related activities should be recognized in the national maritime spatial plans to facilitate long-term planning.

• Bansfield, Danielle; Spilling, Kristian; Mikola, Anna & Piiparinen, Jonna (2021). Bioflocculation of Euglena gracilis via direct application of fungal filaments: a rapid harvesting method. Journal of Applied Phycology. ISSN 0921-8971. 34, p. 321–334. doi: 10.1007/s10811-021-02651-5. Full text in Research Archive
• Baumann, Moritz; Taucher, Jan; Paul, Allanah J.; Heinemann, Malte; Vanharanta, Mari & Bach, Lennart T. [Show all 12 contributors for this article] (2021). Effect of Intensity and Mode of Artificial Upwelling on Particle Flux and Carbon Export. Frontiers in Marine Science. ISSN 2296-7745. 8. doi: 10.3389/fmars.2021.742142. Full text in Research Archive
• Spilling, Kristian; Seppälä, Jukka; Schwenk, Dagmar; Rischer, Heiko & Tamminen, Timo (2021). Variation in the fatty acid profiles of two cold water diatoms grown under different temperature, light, and nutrient regimes. Journal of Applied Phycology. ISSN 0921-8971. 33(3), p. 1447–1455. doi: 10.1007/s10811-021-02380-9.
• Camarena-Gómez, María Teresa; Ruiz-González, Clara; Piiparinen, Jonna; Lipsewers, Tobias; Sobrino, Cristina & Logares, Ramiro [Show all 7 contributors for this article] (2021). Bacterioplankton dynamics driven by interannual and spatial variation in diatom and dinoflagellate spring bloom communities in the Baltic Sea. Limnology and Oceanography. ISSN 0024-3590. 66(1), p. 255–271. doi: 10.1002/lno.11601. Full text in Research Archive
• Lähteenmäki-Uutela, Anu; Rahikainen, Moona; Camarena-Gómez, María Teresa; Piiparinen, Jonna; Spilling, Kristian & Yang, Baoru (2021). European Union legislation on macroalgae products. Aquaculture International. ISSN 0967-6120. 29, p. 487–509. doi: 10.1007/s10499-020-00633-x. Full text in Research Archive
• Bach, Lennart Thomas; Paul, Allanah Joy; Boxhammer, Tim; von der Esch, Elisabeth; Graco, Michelle & Schulz, Kai Georg [Show all 46 contributors for this article] (2020). Factors controlling plankton community production, export flux, and particulate matter stoichiometry in the coastal upwelling system off Peru. Biogeosciences. ISSN 1726-4170. 17(19), p. 4831–4852. doi: 10.5194/bg-17-4831-2020.
• Piwosz, Kasia; Vrdoljak, Ana; Frenken, Thijs; Gonzalez-Olalla, Juan Manuel; Šantić, Danijela & McKay, R. Michael [Show all 17 contributors for this article] (2020). Erratum for Piwosz et al., “Light and Primary Production Shape Bacterial Activity and Community Composition of Aerobic Anoxygenic Phototrophic Bacteria in a Microcosm Experiment”. mSphere. ISSN 2379-5042. 5(4). doi: 10.1128/mSphere.00673-20.
• Piwosz, Kasia; Vrdoljak, Ana; Frenken, Thijs; González-Olalla, Juan Manuel; Šantić, Danijela & McKay, R. Michael [Show all 17 contributors for this article] (2020). Light and primary production shape bacterial activity and community composition of aerobic anoxygenic phototrophic bacteria in a microcosm experiment. mSphere. ISSN 2379-5042. 5(4). doi: 10.1128/MSPHERE.00354-20. Full text in Research Archive

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• Spilling, Kristian; Asmala, Eero; Haavisto, Noora; Haraguchi, Lumi; Kraft, Kaisa & Lehto, Anne-Mari [Show all 14 contributors for this article] (2023). Corrigendum to “Brownification affects phytoplankton community composition but not primary productivity in eutrophic coastal waters: A mesocosm experiment in the Baltic Sea” [Sci. Total Environ. 841 (2022): 156510] (Science of the Total Environment (2022) 841, (S0048969722036075), (10.1016/j.scitotenv.2022.156510)). Science of the Total Environment. ISSN 0048-9697. 871. doi: 10.1016/j.scitotenv.2023.162040.
• Spilling, Kristian; Tedesco, Letizia; Klais, Riina & Olli, Kalle (2019). Editorial: Changing plankton communities: Causes, effects and consequences. Frontiers in Marine Science. ISSN 2296-7745. 6(May). doi: 10.3389/fmars.2019.00272.

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