Abstract
Deep-sea sponge grounds are habitat-forming benthic communities characterized by high biomass and structural complexity. Despite their ecological significance, their role for the deep-sea environment remains poorly understood and their functioning is often inferred from ex situ studies. We hypothesized that deep-sea sponge grounds exhibit substantially higher respiration and nutrient turnover than surrounding soft sediments, making them hotspots of carbon and nutrient cycling in the deep sea. Integrated respiration and nutrient cycling were quantified in a sponge ground on the summit of an Arctic seamount (Schulz Bank, ~ 580 m depth). We used in-situ incubation chambers measuring oxygen consumption, prokaryotic cell removal, and inorganic nutrient fluxes. Respiration rates ranged from 0.13 to 0.93 mmol O₂ m⁻² h⁻¹, which is comparable to cold-water coral reefs and up to 7–21 times higher than reported for soft sediments of the Arctic deep sea. This indicates a high organic carbon demand exceeding surface-derived supply, suggesting the uptake of additional food resources. All incubations showed net release of ammonium, phosphate, nitrite and nitrate, with fluxes correlating with sponge biomass. Our results demonstrate that deep-sea sponge grounds function as hotspots of carbon and nutrient cycling and suggest distinct functional contributions of sponge groups and their microbiome.
Data availability
The data supporting this study are available from the corresponding author upon request.
References
Klitgaard, A. B. & Tendal, O. S. Distribution and species composition of mass occurrences of large-sized sponges in the northeast Atlantic. Prog Oceanogr. 61, 57–98 (2004).
Roberts, J. M., Wheeler, A. J. & Freiwald, A. Reefs of the Deep: The Biology and Geology of Cold-Water Coral Ecosystems. Science 312, 543–547 (2006).
Bart, M. C., Hudspith, M., Rapp, H. T., Verdonschot, P. F. M. & de Goeij, J. M. A Deep-Sea Sponge Loop? Sponges Transfer Dissolved and Particulate Organic Carbon and Nitrogen to Associated Fauna. Front Mar. Sci 8, 604879 (2021).
Hanz, U. et al. The important role of sponges in carbon and nitrogen cycling in a deep-sea biological hotspot. Funct. Ecol. 36, 2188–2199 (2022).
Maldonado, M. et al. Massive silicon utilization facilitated by a benthic-pelagic coupled feedback sustains deep-sea sponge aggregations. Limnol. Oceanogr. 66, 366–391 (2021).
Roberts, E. M. et al. Occurrence records of Geodia species (Porifera, Demospongiae, Tetractinellida) from the North Atlantic and Arctic Oceans. PANGAEA (2021). https://doi.org/10.1594/PANGAEA.924531
Hawkes, N. et al. Glass sponge grounds on the Scotian Shelf and their associated biodiversity. Mar. Ecol. Prog Ser. 614, 91–109 (2019).
Vafeiadou, A., Fragkopoulou, E. & Assis, J. A global dataset of demosponge distribution records. Data Brief. 53, 110200 (2024).
Liu, F. et al. Can Environmental Conditions at North Atlantic Deep-Sea Habitats Be Predicted Several Years Ahead? ——Taking Sponge Habitats as an Example. Front Mar. Sci 8, 703297 (2021).
Tabachnick, K. R., Van Soest, R., van Kempen Th, M. & Braekamn, J. Distribution of recent Hexactinellida in Sponges in Time and Space(eds van Soest, R. et al.) 225–232 (Balkema, Rotterdam, 1994).
Maldonado, M. et al. Sponge grounds as key marine habitats: a synthetic review of types, structure, functional roles, and conservation concerns inMarine Animal Forests(eds Rossi, S. et al.) (Springer, Cham, 2017).
Hogg, M. M. et al. Deep-sea sponge grounds: reservoirs of biodiversity. UNEP-WCMC Biodivers. Ser. 32, 1–86 (2010).
Meyer, H. K., Roberts, E. M., Rapp, H. T. & Davies, A. J. Spatial patterns of arctic sponge ground fauna and demersal fish are detectable in autonomous underwater vehicle (AUV) imagery. Deep Sea Res. Part. Oceanogr. Res. Pap. 153, 103137 (2019).
Weisz, J. B., Lindquist, N. & Martens, C. S. Do associated microbial abundances impact marine demosponge pumping rates and tissue densities? Oecologia 155, 367–376 (2008).
Yahel, G., Whitney, F., Reiswig, H. M., Eerkes-Medrano, D. I. & Leys, S. P. In situ feeding and metabolism of glass sponges (Hexactinellida, Porifera) studied in a deep temperate fjord with a remotely operated submersible. Limnol. Oceanogr. 52, 428–440 (2007).
Morganti, T. M., Ribes, M., Yahel, G. & Coma, R. Size Is the Major Determinant of Pumping Rates in Marine Sponges. Front. Physiol. 10, 1474 (2019).
Kahn, A. S., Yahel, G., Chu, J. W. F., Tunnicliffe, V. & Leys, S. P. Benthic grazing and carbon sequestration by deep-water glass sponge reefs: Deep-water glass sponge reefs. Limnol. Oceanogr. 60, 78–88 (2015).
Leys, S. P., Kahn, A. S., Fang, J. K. H., Kutti, T. & Bannister, R. J. Phagocytosis of microbial symbionts balances the carbon and nitrogen budget for the deep-water boreal sponge Geodia barretti. Limnol. Oceanogr. 63, 187–202 (2018).
Hansell, D. A., Carlson, C. A., Repeta, D. J. & Schlitzer, R. Dissolved organic matter in the ocean: A controversy stimulates new insights. Oceanography 22, 202–211 (2009).
Bart, M. C. et al. Dissolved organic carbon (DOC) is essential to balance the metabolic demands of four dominant North-Atlantic deep‐sea sponges. Limnol. Oceanogr. 66, 925–938 (2021).
Bart, M. C. et al. Differential processing of dissolved and particulate organic matter by deep-sea sponges and their microbial symbionts. Sci. Rep. 10, 17515 (2020).
Olinger, L. K., Strangman, W. K., McMurray, S. E. & Pawlik, J. R. Sponges With Microbial Symbionts Transform Dissolved Organic Matter and Take Up Organohalides. Front Mar. Sci 8, 665789 (2021).
Maier, S. R. et al. Recycling pathways in cold-water coral reefs: Use of dissolved organic matter and bacteria by key suspension feeding taxa. Sci. Rep. 10, 9942 (2020).
Rix, L. et al. Coral mucus fuels the sponge loop in warm-and cold-water coral reef ecosystems. Sci. Rep. 6, 18715 (2016).
Taylor, M. W., Radax, R., Steger, D. & Wagner, M. Sponge-Associated Microorganisms: Evolution, Ecology, and Biotechnological Potential. Microbiol. Mol. Biol. Rev. 71, 295–347 (2007).
Maldonado, M. et al. A Microbial Nitrogen Engine Modulated by Bacteriosyncytia in Hexactinellid Sponges: Ecological Implications for Deep-Sea Communities. Front Mar. Sci 8, 638505 (2021).
Busch, K. et al. Biodiversity, environmental drivers, and sustainability of the global deep-sea sponge microbiome. Nat. Commun. 13, 5160 (2022).
Hentschel, U. et al. Microbial Diversity of Marine Sponges. in Sponges (Porifera) (ed Müller, W. E. G.) vol. 37 59–88 (Springer Berlin Heidelberg, Berlin, Heidelberg, (2003).
De Kluijver, A. et al. An integrative model of carbon and nitrogen metabolism in a common deep-sea sponge (Geodia barretti). Front. Mar. Sci. 7, 596251 (2021).
Garritano, A. N. et al. Species-specific relationships between deep sea sponges and their symbiotic Nitrosopumilaceae. ISME J. 17, 1517–1519 (2023).
Radax, R. et al. Metatranscriptomics of the marine sponge Geodia barretti: tackling phylogeny and function of its microbial community. Environ. Microbiol. 14, 1308–1324 (2012).
Hudspith, M. et al. Quantifying sponge host and microbial symbiont contribution to dissolved organic matter uptake through cell separation. Mar. Ecol. Prog Ser. 670, 1–13 (2021).
Rix, L. et al. Heterotrophy in the earliest gut: a single-cell view of heterotrophic carbon and nitrogen assimilation in sponge-microbe symbioses. ISME J. 14, 2554–2567 (2020).
Meyer, H. K. et al. Beyond the tip of the seamount: Distinct megabenthic communities found beyond the charismatic summit sponge ground on an arctic seamount (Schulz Bank, Arctic Mid-Ocean Ridge). Deep Sea Res. Part. Oceanogr. Res. Pap. 191, 103920 (2023).
Hanz, U. et al. Long-term Observations Reveal Environmental Conditions and Food Supply Mechanisms at an Arctic Deep-Sea Sponge Ground. J. Geophys. Res. Oceans. 126, e2020JC016776 (2021).
Roberts, E. M. et al. Oceanographic setting and short-timescale environmental variability at an Arctic seamount sponge ground. Deep Sea Res. Part. Oceanogr. Res. Pap. 138, 98–113 (2018).
Stratmann, T., Mevenkamp, L., Sweetman, A. K., Vanreusel, A. & Van Oevelen, D. Has phytodetritus processing by an abyssal soft-sediment community recovered 26 years after an experimental disturbance? Front. Mar. Sci. 5, 59 (2018).
Brussaard, C. P. D. Optimization of Procedures for Counting Viruses by Flow Cytometry. Appl. Environ. Microbiol. 70, 1506–1513 (2004).
Yahel, G., Sharp, J. H., Marie, D., Häse2, C. & Genin, A. In situ feeding and element removal in the symbiont-bearing sponge Theonella swinhoei: Bulk DOC is the major source for carbon. Limnol. Oceanogr. 48, 141–149 (2003).
Cathalot, C. et al. Cold-water coral reefs and adjacent sponge grounds: hotspots of benthic respiration and organic carbon cycling in the deep sea. Front. Mar. Sci. 2, 37 (2015).
Tanioka, T. & Matsumoto, K. Stability of Marine Organic Matter Respiration Stoichiometry. Geophys Res. Lett 47 (1), e2019GL085564 (2020).
Fukuda, R., Ogawa, H., Nagata, T. & Koike, I. Direct Determination of Carbon and Nitrogen Contents of Natural Bacterial Assemblages in Marine Environments. Appl. Environ. Microbiol. 64, 3352–3358 (1998).
Donis, D., McGinnis, D. F., Holtappels, M., Felden, J. & Wenzhoefer, F. Assessing benthic oxygen fluxes in oligotrophic deep sea sediments (HAUSGARTEN observatory). Deep Sea Res. Part. Oceanogr. Res. Pap. 111, 1–10 (2016).
Kutti, T., Bannister, R. J. & Fosså, J. H. Community structure and ecological function of deep-water sponge grounds in the Traenadypet MPA—Northern Norwegian continental shelf. Cont. Shelf Res. 69, 21–30 (2013).
Wurz, E. et al. The Hexactinellid Deep-Water Sponge Vazella pourtalesii (Schmidt, 1870) (Rossellidae) Copes With Temporarily Elevated Concentrations of Suspended Natural Sediment. Front Mar. Sci 8, 611539 (2021).
van Oevelen, D. et al. The cold-water coral community as hotspot of carbon cycling on continental margins: A food-web analysis from Rockall Bank (northeast Atlantic). Limnol. Oceanogr. 54, 1829–1844 (2009).
De Froe, E. et al. Benthic oxygen and nitrogen exchange on a cold-water coral reef in the North-East Atlantic Ocean. Front. Mar. Sci. 6, 665 (2019).
Rovelli, L. et al. Benthic O2 uptake of two cold-water coral communities estimated with the non-invasive eddy correlation technique. Mar. Ecol. Prog Ser. 525, 97–104 (2015).
Maier, S. R. et al. On the paradox of thriving cold-water coral reefs in the food-limited deep sea. Biol. Rev. 98, 1768–1795 (2023).
Andersson, J. H. et al. Respiration patterns in the deep ocean. Geophys Res. Lett 31, L03304 (2004).
Ramirez-Llodra, E. et al. The emerging picture of a diverse deep Arctic Ocean seafloor: From habitats to ecosystems. Elem. Sci. Anthr. 12, 00140 (2024).
Stratmann, T., Soetaert, K., Wei, C. L., Lin, Y. S. & Van Oevelen, D. The SCOC database, a large, open, and global database with sediment community oxygen consumption rates. Sci. Data. 6, 242 (2019).
Smith, C. R., De Leo, F. C., Bernardino, A. F., Sweetman, A. K. & Arbizu, P. M. Abyssal food limitation, ecosystem structure and climate change. Trends Ecol. Evol. 23, 518–528 (2008).
Mueller, B. et al. Natural Diet of Coral-Excavating Sponges Consists Mainly of Dissolved Organic Carbon (DOC). PLOS ONE. 9, e90152 (2014).
de Goeij, J. M., Berg, H., van van den, Oostveen, M. M., Epping, E. H. G. & van Duyl, F. C. Major bulk dissolved organic carbon (DOC) removal by encrusting coral reef cavity sponges. Mar. Ecol. Prog Ser. 357, 139–151 (2008).
Hanz, U. et al. Seasonal Variability in Near-bed Environmental Conditions in the Vazella pourtalesii Glass Sponge Grounds of the Scotian Shelf. Front Mar. Sci 7, 597682 (2021).
Ward, B. B. et al. Ammonia-oxidizing bacterial community composition in estuarine and oceanic environments assessed using a functional gene microarray. Environ. Microbiol. 9, 2522–2538 (2007).
Tang, W. et al. Database of nitrification and nitrifiers in the global ocean. Earth Syst. Sci. Data. 15, 5039–5077 (2023).
Pachiadaki, M. G. et al. Major role of nitrite-oxidizing bacteria in dark ocean carbon fixation. Science 358, 1046–1051 (2017).
Hoffmann, F. et al. Complex nitrogen cycling in the sponge Geodia barretti. Environ. Microbiol. 11, 2228–2243 (2009).
Kumala, L. & Canfield, D. E. Contraction Dynamics and Respiration of Small Single-Osculum Explants of the Demosponge Halichondria panicea. Front Mar. Sci 5, 410 (2018).
Stratmann, T. et al. Nutrient fluxes, oxygen consumption and fatty acid composition from deep-water demo- and hexactinellid sponges from New Zealand. Deep Sea Res. Part. Oceanogr. Res. Pap. 214, 104416 (2024).
Maldonado, M., Bayer, K. & López-Acosta, M. Nitrogen and Phosphorus Cycling Through Marine Sponges: Physiology, Cytology, Genomics, and Ecological Implications. in Frontiers in Invertebrate Physiology: A Collection of Reviews (Apple Academic, 2024).
Godwin, C. M. & Cotner, J. B. Aquatic heterotrophic bacteria have highly flexible phosphorus content and biomass stoichiometry. ISME J. 9, 2324–2327 (2015).
Thamdrup, B. & Dalsgaard, T. Production of N2 through Anaerobic Ammonium Oxidation Coupled to Nitrate Reduction in Marine Sediments. Am. Soc. Microbiol. https://doi.org/10.1128/AEM.68.3.1312 (2002). -1318.2002.
De Goeij, J. M. et al. Surviving in a Marine Desert: The Sponge Loop Retains Resources Within Coral Reefs. Science 342, 108–110 (2013).
Busch, K. et al. On giant shoulders: how a seamount affects the microbial community composition of seawater and sponges. Biogeosciences 17, 3471–3486 (2020).
Chai, G., Li, J. & Li, Z. The interactive effects of ocean acidification and warming on bioeroding sponge Spheciospongia vesparium microbiome indicated by metatranscriptomics. Microbiol. Res. 278, 127542 (2024).
Maggioni, F. et al. Warm-adapted sponges resist thermal stress by reallocating carbon and nitrogen resources from cell turnover to somatic growth. Limnol. Oceanogr. 69, 976–991 (2024).
Strand, R. et al. The response of a boreal deep-sea sponge holobiont to acute thermal stress. Sci. Rep. 7, 1660 (2017).
Liu, W., Fedorov, A. V., Xie, S. P. & Hu, S. Climate impacts of a weakened Atlantic Meridional Overturning Circulation in a warming climate. Sci. Adv. 6, eaaz4876 (2020).
Sanni, S. et al. Environmental risk framework and research recommendations for SMS mining in the Norwegian Arctic mid-ocean ridge. Front Mar. Sci 12, 1651042 (2025).
Wurz, E. et al. Adverse effects of crushed seafloor massive sulphide deposits on the boreal deep-sea sponge Geodia barretti Bowerbank, 1858 and its associated fauna. Deep Sea Res. Part. Oceanogr. Res. Pap. 208, 104311 (2024).
Acknowledgements
We thank the crews and scientific parties of RV G. O. Sars cruise GS2017110 for their technical support. We thank Stig Vågenes and his ROV Ægir 6000 team for the exceptional skill and precision during the dives.
Funding
Open Access funding enabled and organized by Projekt DEAL. This research has been performed in the scope of the SponGES project, which received funding from the European Union’s Horizon 2020 research and innovation program under grant agreement No. 679849. I acknowledge support by the Open Access publication fund of Alfred-Wegener-Institut Helmholtz- Zentrum für Polar- und Meeresforschung. Open Access funding enabled and organized by Projekt DEAL.
Author information
Authors and Affiliations
Contributions
UH, BM, MB, JMG, FM designed the study. UH, BM, MB, JMG, FM, HTR participated in sampling. UH, BM, MB, JMG, FM conducted the incubations. KB did microbial analyses. HTR conducted sponge taxonomic analysis. UH performed the data analysis and visualization. UH wrote the manuscript. UH, BM, MB, JMG, FM, KB, GJR discussed the results, reviewed and edited the manuscript.
Corresponding author
Ethics declarations
Competing interests
The authors declare no competing interests.
Additional information
Publisher’s note
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Supplementary Information
Below is the link to the electronic supplementary material.
Rights and permissions
Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.
About this article
Cite this article
Hanz, U., Mueller, B., Bart, M.C. et al. Unveiling in situ oxygen, carbon and nutrient cycling of a sponge-driven biological hotspot in the arctic. Sci Rep (2026). https://doi.org/10.1038/s41598-026-41798-4
Received:
Accepted:
Published:
DOI: https://doi.org/10.1038/s41598-026-41798-4