7 - Litteratur
Bradshaw, C, Tjensvoll, I., Sköld, M., Allan, I.J., Molvaer, J., Magnusson, J., Naes, K. and Nilsson, H.C. (2012). Bottom trawling resuspends sediment and releases bioavailable contaminants in a polluted fjord, Environmental Pollution, 170, 232-241. https://doi.org/10.1016/j.envpol.2012.06.019.
Buhl-Mortensen, P. (2017). Coral reefs in the Southern Barents Sea: habitat description and the effects of bottom fishing. Marine Biology Research, 13(10), 1027-1040.
Buhl-Mortensen, L., Aglen, A., Breen, M., Buhl-Mortensen, P., Ervik, A., Husa, V., et al. (2013). Impacts of Fisheries and Aquaculture on Sediments and Benthic Fauna: Suggestions for New Management Approaches. Fisken og Havet.
Buhl-Mortensen, L., Ellingsen, K.E., Buhl-Mortensen, P., Skaar, K.L. and Gonzales-Mirelis, G. (2015). Trawling disturbance on megabenthos and sediment in the Barents Sea: chronic effects on density, diversity, and composition. ICES Journal of Marine Science. Doi:10.1093/icesjms/fsv200
Certain, G., Jørgensen, L.L., Christel, I., Planque, B. and Vinceny, B. (2015). Mapping the vulnerability of animal community to pressure in marine systems: Disentangling impact types and integrating their effect from the individual to the community level. ICES Journal of Marine Science. Doi:10.1093/icesjms/fsv003
Desprez M. (2000). Physical and biological impact of marine aggregate extraction along the French coast of the Eastern English Channel: short- and long-term post-dredging restoration. ICES Journal of Marine Science, 57,1428-1438
Diesing, M., Thorsnes, T. and Bjarnadottir, L.R. (2021). Organic carbon densities and accumulation rates in surface sediments of the North Sea and Skagerrak. Biogeosciences, 18:2139–2160.
Durrieu de Madron, X., Ferre, B., Le Corre, G., Grenz, C., Conan, P., Pujo-Pay, M., Buscail, R. and Bodiot, O. (2005). Trawling-induced resuspension and dispersal of muddy sediments and dissolved elements in the Gulf of Lion (NW Mediterranean). Continental Shelf Research. 25, 2387-2409.
Eigaard, O. R., Bastardie, F., Breen, M., Dinesen, G. E., Hintzen, N. T., Laffargue, P., Mortensen, L.O. et al. (2016). Estimating seabed pressure from demersal trawls, seines, and dredges based on gear design and dimensions. ICES Journal of Marine Science, doi:10.1093/icesjms/fsv099
Eigaard, O. R., Bastardie, F., Hintzen, N. T., Buhl-Mortensen, L., Buhl-Mortensen., P., Catarino, R., Dinesen, G. E. et al. (2017). The footprint of bottom trawling in European waters: distribution, intensity, and seabed integrity. ICES Journal of Marine Science, doi:10.1093/icesjms/fsw194
Epstein, G., Middelburg, J. J., Hawkins, J. P., Norris, C. R. and Roberts, C. M. (2022). The impact of mobile demersal fishing on carbon storage in seabed sediments. Global Change Biology, 28, 2875– 2894. https://doi.org/10.1111/gcb.16105
Eriksen, E., van der Meeren, G.I., Nilsen, B.M., von Quillfeldt, C.H. og Johnsen, H. (2021). Særlig verdifulle og sårbare områder (SVO) i norske havområder – Miljøverdi. Rapport fra Havforskningen, Nr. 2021-26.
Fosså, J.H., Kutti, T., Buhl-Mortensen, P. og Skjoldal, H.R. (2015). Vurdering av norske korallrev. Rapport fra Havforskningen, Nr. 8-2015.
Freese, L., Auster, P.J., Heifetz, J. and Wing, B.L. (1999). Effects of trawling on seafloor habitat and associated invertebrate taxa in the Gulf of Alaska. Marine Ecology Progress Series, 182, 119-126.
Hansson, M., Lindegarth, M., Valentinsson, D. and Ulmestrand, M. (2000). Effects of shrimp-trawling on abundance of benthic macrofauna in Gullmarsfjorden, Sweden. Marine Ecology Progress Series, 198, 191-201.
He, P., Suuronen, P., Ferro., R.S.T. and Lansley, J. (2021). Classification and illustrated definition of fishing gears. FAO Fisheries and Aquaculture Technical Paper No. 672. Rome, FAO. https://doi.org/10.4060/cb4966en
Hiddink, J.G., Jennings, S., Sciberras, M., Szostek, C.L., Hughes, K.M., Ellis, N., Rijnsdorp, A.D. et al. (2017). Global analysis of depletion and recovery of seabed biota after bottom trawling disturbance. Proceedings of the National Academy of Science USA, 114(31), 8301-8306.
Hilborn, R. and Kaiser, M.J. (2022). A path forward for analysing the impacts of marine protected areas. Nature. https://doi.org/10.1038/s41586-022-04775-1
Humborstad, O.-B., Jørgensen, T. and Grotmol, S. (2006). Exposure of cod Gadus morhua to resuspended sediment: an experimental study of impact of bottom trawling. Marine Ecology Progress Series, 309, 247-254.
Humborstad, O.-B., Nøttestad, L., Løkkeborg, S. and Rapp, H.T. (2004). RoxAnn bottom classification system, sidescan sonar and video-sledge: Spatial resolution and their use in assessing trawling impacts. ICES Journal of Marine Science, 61, 53-63.
ICES. 2021. Working Group on the Integrated Assessments of the Barents Sea (WGIBAR). ICES Scientific Reports. 3:77. 236 pp. https://doi.org/10.17895/ices.pub.8241
ICES. 2022. Working Group on the Integrated Assessments of the Barents Sea (WGIBAR). ICES Scientific Reports. 4:50. 235 pp. http://doi.org/10.17895/ices.pub.20051438
Jennings, S., Pinnegar, J.K., Polunin, N.V.C. and Warr, K.J. (2001). Impacts of trawling disturbance on the trophic structure of benthic invertebrate communities. Marine Ecology Progress Series, 213, 127-142.
Jørgensen, L.L., Bakke, G. and Hoel, A.H. (2020). Responding to global warming: New fisheries management measures in the Arctic. Progress in Oceanography. https://doi.org/10.1016/j.pocean.2020.102423
Jørgensen, L.L., Ljubin, P., Skjoldal, H.R., Ingvaldsen, R.B., Anisimova, N. and Manushin, I. (2015). Distribution of benthic megafauna in the Barents Sea: baseline for an ecosystem approach to management. ICES Journal of Marine Science, 72 (2), 595-613.
Jørgensen, L.L., Planque, B., Thangstad, T.H. and Certain, G. (2016). Vulnerability of megabenthic species to trawling in the Barents Sea . ICES Journal of Marine Science. DOI: 10.1093/icesjms/fsv107 .
Jørgensen, L.L., Primicerio, R., Ingvaldsen, R.B., Fossheim, M., Strelkova, N., Thangstad, T.H., Manushin, I. and Zakharov, D. (2019). Impact of multiple stressors on seabed fauna in a warming Arctic. Marine Ecology Progress Series, 608, 1-12.
Kaiser, M. J. , Clarke, K. D. , Hinz, H. , Austen, M. C. V. , Somerfield, P. J. , and Karakassis, I. ( 2006 ). Global analysis of response and recovery of benthic biota to fishing . Marine Ecology Progress Series, 311 , 1- 14 .
Kędra, M., Renaud, P. E., and Andrade, H. (2017). Epibenthic diversity and productivity on a heavily trawled Barents Sea bank (Tromsøflaket). Oceanologia, 59(2), 93-101.
Kenchington, E.L.R., Gilkinson, K.D., MacIssac, K.C., Bourbonnais-Boyce, C., Kenchington, T.J., Smith, S.J. and Gordon Jr., D.C. (2006). Effects of experimental otter trawling on benthic assemblages on Western Bank, northwest Atlantic Ocean. Journal of Sea Research, 56, 249-270.
Kenchington, E.L.R., Prena, J., Gilkinson, K.D., Gordon Jr., D.C., MacIssac, K., Bourbonnais, C., Schwinghamer, P.J., Rowell, T.W., McKeown, D.L. and Vass, W.P. 2001. Effects of experimental otter trawling on the macrofauna of a sandy bottom ecosystem on the Grand Banks of Newfoundland. Canadian Journal of Fisheries and Aquatic Science, 58, 1043-1057.
Kroodsma, D.A., Mayorga, J., Hochberg, T. et al. (2018). Tracking the global footprint of fisheries. Science, 359,904–908. https://doi.org/10.1126/science.aao5646
Kutti, T., Høisæter, T., Rapp. H.T., Humborstad, O.-B., Løkkeborg, S. and Nøttestad, L. (2005). Immediate effects of experimental otter trawling on a sub-artic benthic assemblage inside the Bear Island Fishery Protection Zone in the Barents Sea. American Fishery Society Symposia, 41: 519-528.
Lindegarth, M., Valentinsson, D., Hansson, M. and Ulmestrand, M. 2000. Interpreting large-scale experiments on effects of trawling on benthic fauna: an empirical test of the potential effects of spatial confounding in experiments without replicated control and trawled areas. Journal of Experimental Marine Biology and Ecology, 245, 155-169.
Lucchetti, A. and Sala, A. (2012). Impact and performance of Mediterranean fishing gear by side-scan sonar technology. Canadian Journal of Fisheries and Aquatic Science, 69, 1806-1816.
Lyubin, P. A., Anisimova, A. A. and Manushin, I. E. (2011). Long-term effects on benthos of the use of bottom fishing gears. In: Jakobsen, T. and Ozhigin, V.K. (Eds.), The Barents Sea. Ecosystem, Resources, Management, 768–775, Tapir Academic Press, Trondheim.
Løkkeborg, S. (2005). Impacts of trawling and scallop dredging on benthic habitats and communities.
FAO Fisheries Technical Paper. No. 472. Rome, FAO. 2005. 58p.
Løkkeborg, S. and Fosså, J.H. (2011). Impacts of bottom trawling on benthic habitats. In: Jakobsen, T. and Ozhigin, V.K. (Eds.), The Barents Sea. Ecosystem, Resources, Management, pp. 760-767, Tapir Academic Press, Trondheim.
MarLIN, 2006. BIOTIC - Biological Traits Information Catalogue. Marine Life Information Network. Plymouth: Marine Biological Association of the United Kingdom. Available from www.marlin.ac.uk/biotic
Martín, J., Puig, P., Masque, P., Palanques, A., & Sanchez-Gomez, A. (2014a). Impact of bottom trawling on deep-sea sediment properties along the flanks of a submarine canyon. PLoS One, 9(8), e104536. https://doi.org/10.1371/journal.pone.0104536
Mazor, T., Pitcher, C.R., Rochester, W., Kaiser, M.J., Hiddink, J.G., Jennings, S., Amoroso, R. et al. (2020). Trawl fishing impacts on the status of seabed fauna in diverse regions of the globe. Fish and Fisheries, 22, 72-86.
McConnaughey, R.A. and Syrjala, S.E. (2014). Short-term effects of bottom trawling and a storm event on soft-bottom benthos in the Bering Sea. ICES Journal of Marine Science, 71, 2469-2483.
Mengual, B., Cayocca, F., Hir, P.L., Draye, R., Laffargue, P., Vincent, B. and Garlan, T. (2016). Influence of bottom trawling on sediment resuspension in the “Grande-Vasiere” area (Bay of Biscay, France. Ocean Dynamics, 66, 1181-1207.
de Moura Neves, B., Edinger, E. and Hayes, V.W. (2018). Morphology and composition of the internal axis in two morphologically contrasting deep-water sea pens (Cnidaria:Octocorallia). Journal of Natural History, 52, 659-685.
Oberle F. K. J., Storlazzi, C. D., & Hanebuth, T. J. J. (2016). What a drag: Quantifying the global impact of chronic bottom trawling on continental shelf sediment. Journal of Marine Systems, 159, 109–119. https://doi.org/10.1016/j.jmarsys.2015.12.007
O`Neill, F.G., Summerbell, K. and Breen, M. (2009). An underwater laser stripe seabed profiler to measure the physical impact of towed gear components on the seabed. Fisheries Research, 99, 234-238.
Pitcher, C.R., Hiddink, J.G., Jennings, S., Collie, J., Parma, A.M., Amoroso, R., Mazor, T., Sciberras, M., McConnaughey, R.A., Rijnsdorp, A.D., Kaiser, M.J., Suuronen, P., Hilborn, R. (2022). Trawl impacts on the relative status of biotic communities of seabed sedimentary habitats in 24 regions worldwide. Proceedings of the National Academy of Science USA. 119(2): e2109449119. https://doi.org/10.1073/pnas.2109449119
Prena, J., Schwinghamer, P., Rowell, T.W., Gordon Jr., D.C., Gilkinson, K.D., Vass, W.P. and McKeown, D.L. 1999. Experimental otter trawling on a sandy bottom ecosystem of the Grand Banks of Newfoundland: analysis of trawl bycatch and effects on epifauna. Marine Ecology Progress Series, 181, 107-124.
Puig, P., Canals, M., Company, J. B., Martin, J., Amblas, D., Lastras, G., Palanques, A. et al. (2012). Ploughing the deep sea floor. Nature, 489, 286-290. https://doi.org/10.1038/nature11410
Pusceddu, A., Bianchelli, S., Martin, J., Puig, P., Palanques, A., Masqué. P. and Danovaro, R. (2014). Chronic and intensive bottom trawling impairs deep-sea biodiversity and ecosystem functioning. Proceedings of the National Acadamy of Science USA, 111, 8861-8866 https://doi.org/10.1073/pnas.1405454111
Ramirez-Llodra, E., Rinde, E., Gundersen, H. et al. (2016). A snap shot of the short-term response of crustaceans to macrophyte detritus in the deep Oslofjord. Sci Rep 6, 23800. https://doi.org/10.1038/srep23800
Rijnsdorp, A.D., Buys, A.M., Storbeck, F. and Visser, E.G. (1998). Micro-scale distribution of beam trawl effort in the southern North Sea between 1993 and 1996 in relation to the trawling frequency of the sea bed and the impact on benthic organisms. ICES Journal of Marine Science, 55, 403-419.
Rijnsdorp, A. D., Hiddink, J. G., van Denderen, P. D., Hintzen, N. T., Eigaard, O. R., Valanko, S., Bastardie, F. et al. (2020). Different bottom trawl fisheries have a differential impact on the status of the North Sea seafloor habitats. ICES Journal of Marine Science, 77, 1772-1786.
Sala, E., Mayorga, J., Bradley, D., Cabral, R. B., Atwood, T.B., Auber, A. et al. (2021). Protecting the global ocean for biodiversity, food and climate. Nature, https://doi.org/10.1038/s41586-021-03371-z
Sarda, R., Pinedo, S., Gremare, A. and Taboada, S. (2000). Changes in the dynamics of shallow sandy-bottom assemblages due to sand extraction in the Catalan Western Mediterranean Sea. ICES Journal of Marine Science, 57,1446-1453.
Schwinghamer, P., Guigné, J.Y. and Siu, W.C. (1996). Quantifying the impact of trawling on benthic habitat structure using high resolution acoustics and chaos theory. Canadian Journal of Fisheries and Aquatic Science, 53, 288–296.
Schwinghamer, P., Gordon Jr., D.C., Rowell, T.W., Prena, J., McKeown, D.L., Sonnichsen, G. and Guigné, J.Y. (1998). Effects of experimental otter trawling on surficial sediment properties of a sandy-bottom ecosystem on the Grand Banks of Newfoundland. Conservation Biology, 12, 1215–1222.
Sciberras, M., Hiddink, J.G., Jennings, S., Szostek, C.L., Hughes, K.M., Kneafsey, B., Clarke, L.J. et al. (2018). Responses of benthic fauna to experimental bottom fishing: A global meta-analysis. Fish and Fisheries, 19, 698-715.
Smith, C.J., Papadopoulou, K.N. and Diliberto, S. (2000). Impact of otter trawling on an eastern Mediterranean commercial trawl fishing ground. ICES Journal of Marine Science, 57, 1340-1351.
Smeaton, C., & Austin, W. E. N. (2022). Quality not quantity: Prioritizing the management of sedimentary organic matter across continental shelf seas. Geophysical Research Letters, 49, e2021GL097481. https://doi.org/10.1029/2021GL097481
Szostek, C.L., Hiddink, J.G., Sciberras, M., Shepperson, J.L., Thompson, S., Hormbrey, S., Caveen, A. et al. (2022). A tool to estimate the contribution of fishing gear modifications to reduce benthic impact. Journal of Industrial Ecology. https://doi.org/10.1111/jiec.13366
Tuck, I.D., Hall, S.J., Robertson, M.R., Armstrong, E. and Basford, D.J. (1998). Effects of physical trawling disturbance in a previously unfished sheltered Scottish sea loch.
Marine Ecology Progress Series, 162, 227–242.
Underwood, A.J. 1992. Beyond BACI: the detection of environmental impacts on populations in the real, but variable, world. Journal of Experimental Marine Biology and Ecology, 161, 145-178.
van Denderen, P.D., Bolam, S.G., Hiddink, J.G., Jennings, S., Kenny, A., Rijnsdorp, A.D. and van Kooten, T. (2015). Similar effects of bottom trawling and natural disturbance on composition and function of benthic communities across habitats. Marine Ecology Progress Series, 541, 31-43.
Wainwright, W S. C., & Hopkinson Jr, C. S. (1997). Effects of sediment resuspension on organic matter processing in coastal environments: a simulation model. Journal of Marine Systems, 11(3–4), 353–368.
8 - Appendiks
The remineralization of organic carbon (OC) is defined by recycling processes in which OC, as dead organisms (Asper, 1987) and/or metabolites (Duursma, 1963), are broken-down to smaller molecules until entering the water column as dissolved inorganic carbon (DIC) (Emerson, 2013). The rate of this process may be increased by sediment disturbance from bottom fishing due to the reduced production of flora and fauna, the loss of fine flocculent material, increased sediment resuspension, mixing and transport, and increased oxygen exposure (Epstein et al., 2021). However, some processes such as reduced faunal bioturbation and community respiration, increased off-shelf transport and increases in primary production from the resuspension of nutrients, also induced by bottom trawling activity may lead to a decrease in net OC remineralisation (Epstein et al., 2021). The interaction between both positive and negative feedback mechanisms, makes it challenging to identify the impact of trawling on net OC remineralization and associated increases in DIC which are likely site specific. (Keil, 2017; Snelgrove et al., 2018; LaRowe et al., 2020; Rühl et al., 2020). In a recent review of 49 studies that measured changes in sediment OC associated with bottom fishing, 61% of studies observed no significate effect, 29% showed a decrease in sediment OC and 10% showed an increase in sediment OC (Epstein et al., 2021).
Despite these complexities, it has been estimated that just the uppermost centimetre of sediment may have lost ~0.06 Gt of OC due to historical trawling on global continental slopes (Paradis et al., 2021). Sala et al. (2021) estimate 1.47 Pg (Gt) of aqueous CO2 emissions in the first year after trawling due to OC remineralization with continuous trawling lending to a decline in emissions and the stabilization of values after nine years at about 40% of the initial value. This is equivalent to 0.58 Pg (Gt) per year globally. More locally, on the UK shelf, bottom fishing is estimated to remineralise up to ~0.002 Gt of OC per year, assuming that all resuspended OC is remineralised. (Luisetti et al., 2019). However, there are many uncertainties, assumptions, and simplifications in these estimations (Epstein et al., 2021; Hilborn and Kaiser, 2022), in part, due to a lack of site-specific understanding of the complex interactions that determine rates of remineralisation.
More recent studies have therefore focussed on the potential vulnerability (Black et al., 2022) and the annual cumulative disturbance of sedimentary OC stores (Epstein and Roberts, 2022) without explicitly estimating OC remineralization rates. Although progress has been made in mapping OC sediment stocks in recent years (Seiter et al., 2004; Diesing et al., 2017; Lee et al., 2019; Luisetti et al., 2019; Atwood et al., 2020; Legge et al., 2020; Smeaton et al., 2021), in Norway, only the North Sea and Skagerrak have been mapped to date (Diesing et al., 2021). Estimates of OC remineralization, accumulation and burial rates are even more limited (Berner, 1982; Burdige, 2007; Keil, 2017; Wilkinson et al., 2018; Luisetti et al., 2019; Legge et al., 2020; Diesing et al., 2021). Natural rates of OC remineralization and storage also show large spatial and temporal variability. In general, continental shelf and sublittoral zone sediments in summer show the highest rates of OC remineralization (Middelburg et al., 1996; Tabuchi et al., 2010; Brin et al., 2015; Xue et al., 2015) with rates decreasing at higher latitudes (Fiedler et al., 2016; Bourgeois et al., 2017; Zhao et al., 2018a) and deeper depths (reviewed by Chen et al., 2022). Therefore, levels of OC remineralization in response to bottom disturbance by trawling are likely site (and perhaps seasonally) specific and depend on complex interactions between local sediment (e.g. grain size and OC content and stability), environmental conditions (e.g. temperature and oxygenation), biology (e.g. production and bioturbation), and hydrology (e.g. sediment mixing and transport).
8.1 - Local sediment
Local sediment structure and chemistry will, in part, modulate the effect of bottom trawling on OC remineralization. The local stability of OC in the sediment is dependent on the OC molecular size, structure (Amon and Benner, 1996; Van Kaam-Peters et al., 1998) and functional groups (Deng et al., 2019; Kleber and Lehmann, 2019) as well as mineral-organic associations that may inhibit the decomposing activation of enzymes and microbes (Tietjen and G. Wetzel, 2003; Zimmerman et al., 2004). Organic matter, of which OC is a major constituent, is an umbrella term that encompasses a wide range of different substances (e.g., amino acids, sugars, lipids, and lignin) from marine and terrestrial sources (Burdige, 2007). These individual substances vary in their biogeochemical reactivity or degradability. Organic matter reactivity can be seen as a continuum from easily degradable and short-lived (labile) to hard to degrade and long-lived (refractory) (LaRowe et al., 2020). Marine organic matter (e.g., phytoplankton debris) is typically labile, while organic matter from terrestrial sources (e.g., plant litter and soil organic matter) can be considered refractory. Organic matter in marine sediments is a mixture of organic substances from various sources and with varying reactivities. Highly reactive labile constituents will be remineralized first, followed by less reactive substances (Stumm-Zollinger, 1968). Therefore, the overall reactivity of organic matter is reduced over time (Berner, 1980). Likewise, remineralization rates decrease over time.
The rate of remineralization might also change when refractory organic matter comes into contact with labile organic matter; this is called priming (e.g., Bianchi, 2011). Whilst the drivers controlling priming remain unclear, the process must be regarded as important: Simply measuring the reactivity of OC in marine sediments may not fully capture the vulnerability of stored OC when disturbance mixes refractory OC with a labile fraction (Graves et al., 2022) and thereby alters its reactivity. A recently published meta-analysis concluded that, overall, priming increased remineralization of stable OC with the addition of labile OC (Sanches et al., 2021). Disturbance of sediment OC by bottom trawling and mixing of refractory and labile OC might therefore potentially lead to enhanced remineralization and loss of OC.
The grain size and structure of sediment is also important. Sandy sediments are usually associated with deeper oxygen penetration, advective pore water transport, higher levels of natural disturbance leading to lower OC contents and higher rates of remineralization when compared to muds (Burdige, 2007; Huettel et al., 2014). The limited effects of trawling on OC remineralisation in eutrophic fine sediment and coarse sediments has also been attributed to low baseline OC contents. Indeed, in a recent review, of the 61% of reported studies that show no significant effect of bottom trawling on sediment OC content there was a clear trend toward sandy study sites (Epstein et al., 2021).
However, repeated fishing activity can also affect the structure of sediments particularly in finer sediments in less hydrologically active environments (Kaiser et al., 2002; Trimmer et al., 2005; Martín et al., 2014a; Oberle et al., 2016b). In less hydrologically active depositional environments, the resuspension of finer sediments from deeper layers by trawling may lead to a redeposited surface layer of fine sediments. (Palanques et al., 2014; Oberle et al., 2016b; Tiano et al., 2020). In more hydrologically active environments the resuspension and loss of fine material due to transport can lead to an increase in coarse material towards the surface (Martín et al., 2014a; b; Palanques et al., 2014; Pusceddu et al., 2014; Mengual et al., 2016; Oberle et al., 2016b; Paradis et al., 2021), with changes in the vertical structure affecting the environmental conditions to which sediments are exposed and so rates of remineralisation.
8.2 - Environmental conditions
Local environmental conditions such and temperature and oxygen level to which sediments are exposed determine the rates of OC remineralisation. Sediment microbial communities and their metabolic kinetics are highly influenced by temperature (Nedwell, 1999; Trevathan-Tackett et al., 2018) affecting their ability to degrade OC (Malinverno and Martinez, 2015; Roussel et al., 2015; Zang et al., 2020). Low OC remineralization rates have been, in part, linked to lower temperatures in the deep-sea (Weston and Joye, 2005; D’Hondt et al., 2015) and at higher latitudes (Fiedler et al., 2016; Bourgeois et al., 2017; Zhao et al., 2018a), while increasing temperatures due to climate change have also been linked to increased remineralization (Yamamoto-Kawai et al., 2009; Qi et al., 2020).
Oxygen levels are also critical in determining levels of OC remineralization (Hinojosa et al., 2014; Nierop et al., 2017; Kurian et al., 2020). For example, low oxygen concentrations in northern Pacific sediments have been shown to decrease OC remineralisation (Seiter et al., 2005; Jessen et al., 2017). Increased oxygen levels in the sediments can increase microbial respiration and remineralization activity (Kristensen et al., 1995; Dauwe et al., 2001; Keil, 2017; van de Velde et al., 2018). Natural resuspension and changes to the vertical structure of sediments due to physical disturbance by hydrodynamics (Brodersen et al., 2019) or bioturbation (Aller and Cochran, 2019) increase the depth of oxygen penetration increasing OC remineralisation compared to less disturbed anoxic environments (Glud, 2008; Donis et al., 2016).
Although oxygen penetration and/or sediment oxygen concentrations due to acute physical disturbance, such as bottom fishing activity (Allen and Clarke, 2007; Tiano et al., 2019; De Borger et al., 2021) are often short lived, compared to chronic processes such as bioturbation, with generally fast re-establishment of sediment oxygen gradients, these can vary from the pre-disturbed state. (Allen and Clarke, 2007; Tiano et al., 2019; De Borger et al., 2021). Also, the re-establishment of sediment oxygen gradients and their stability would depend on the intensity of fishing activity in an area. However, even when aerobic respiration is dominant in sediment, remineralization rate can be limited by local sediment conditions such as low OC and nutrient concentration, and low temperatures (0–4 oC) as observed in the deep-sea (Weston and Joye, 2005; D’Hondt et al., 2015).
8.3 - Biology
The effect of bottom trawling on OC storage/remineralisation and net local changes in DIC is greatly dependent on the local community, its function, and sensitivity to trawling pressure.
8.3.1 - Infauna and bioturbation
Benthic fauna is critical to biogeochemical cycling in sediments (Middelburg, 2018; Snelgrove et al., 2018; LaRowe et al., 2020; Rühl et al., 2020). Trawling reduces the abundance and diversity of sessile epifauna and burrowing infauna, decreasing benthic biomass and production (Jennings et al., 2001, 2002; Kaiser et al., 2002; Queirós et al., 2006; Sciberras et al., 2018; Tiano et al., 2020; Tillin et al., 2006). Bioturbation (reworking of sediment particles; Ekdale et al., 1984) and bio-irrigation (reworking of sediment solutes; Meysman et al., 2006) by benthic invertebrates are critical for global nutrient and carbon cycling. These possesses can increase OC remineralisation by changing the vertical structure of the sediment increasing the concentrations of oxygen and other electron acceptors (e.g. nitrate, metal oxides and sulphate), that promote microbial breakdown of OC (Hulthe et al., 1998; Meysman et al., 2006; Arndt et al., 2013; Keil, 2017; Snelgrove et al., 2018; LaRowe et al., 2020).
In addition, the transportation of labile OC from the surface to deeper layers may stimulate the microbial remineralisation of more refractory OC stored in deeper sediment (van Nugteren et al., 2009; Middelburg, 2018). Conversely, the transportation of OC to deeper sediment could increase its chance of burial and long-term storage (van der Molen et al., 2012; Middelburg, 2018; Snelgrove et al., 2018; Rühl et al., 2020; De Borger et al., 2021) although this is shown to be site specific (van Nugteren et al., 2009; Bengtsson et al., 2018; Riekenberg et al., 2020).
In heavily trawled areas, large long-lived burrowing species that have the largest effect on nutrient cycling (Olsgard et al., 2008) are replaced by small-bodied, opportunistic, motile infauna, and larger, highly vagrant, scavenging macrofauna (Jennings et al., 2001; 2002; Kaiser et al., 2002; 2006; Thrush & Dayton, 2002; Tillin et al., 2006). The loss of fauna and flora that stabilize sediments can also lead to increased sediment transport and remineralisation (Roberts, 2007)
8.3.2 - Phytoplankton
Levels of primary production in the water column is a significantly driver of OC content in sediments due to vertical transport of dead material (Seiter et al., 2004; Turner, 2015; Atwood et al., 2020). Primary production may be stimulated by an increase in nutrients entering the water column following sediment disturbance (Fanning et al., 1982; Falcão et al., 2003; de Madron et al., 2005; Polymenakou et al., 2005; Pusceddu et al., 2015). In the Mediterranean, modelling based on trawling experiments has estimated that sediment disturbance by fishing gear could increase local net annual primary production by 15% leading to increased OC settlement. However, this will depend on the hydrodynamic conditions and both the transport of nutrients up to the euphotic zone and transport of dead material to the seabed. It is also possible that decreases in light due increased turbidity from resuspended particles may limit photosynthesis and primary production (Ruffin, 1998; Palanques et al., 2001; Adriano et al., 2005; Cloern et al., 2014; Capuzzo et al., 2015).
8.3.3 - Benthic Algae
Ephemeral macroalgae and microphytobenthos have been shown to recover quickly to trawling disturbance, depending on event frequency (MacIntyre et al., 1996; Ordines et al., 2017). However more sensitive habitats like kelp can take years to recover and coralline algae can require decades to recover (e.g. Dayton et al., 1992; Fragkopoulou et al., 2021). The accumulation rate and stability of sediments, critical OC burial and storage (Middelburg, 2018; LaRowe et al., 2020), is enhanced by benthic micro- and macroalgae (Yallop et al., 1994; Miller et al., 1996; Montserrat et al., 2008). It is possible that in some areas the loss of benthic algae could decrease sediment stability, increased benthic mixing and increased oxygen penetration (even in the absence of repeated trawling) leaded to increased OC remineralisation. In addition to increased DIC realised from OC remineralisation, net increases in local DIC may be confounded by reduced photosynthesis. However, this would depend on the local substrate as many kelp areas in Norway are more associated with rocky bottoms with high rates of sediment transport and low OC accumulation, in addition to being at less risk from trawling.
8.4 - Hydrodynamic activity
The interaction of unidirectional currents and oscillating flows caused by sea surface waves with the seabed shapes the general patterns of sediment distribution across continental shelves. Fine-grained deposits (clays, silts, and muds) tend to accumulate in hydrodynamically quiet settings, e.g. in deep basins, which cannot be reached by wave activity and where current speeds are low. Conversely, coarse-grained deposits (sands and gravels) tend to dominate in hydrodynamically active areas, e.g. in the coastal zone, on shelf banks and the shelf break.
An important parameter that influences the way sediments process OC is permeability, or the resistance to flow of water through the sediment (Bear, 1972). Permeability is loosely correlated with the grain-size of the sediment (Wilson et al., 2008); coarser-grained sediments tend to have higher permeabilities and vice versa. Permeability begins to influence biogeochemical processes in the surface layer of marine sediments when pressure gradients in the benthic environment can drive pore-water flows that transport solutes and small particles more effectively than Brownian molecular motion (Huettel et al., 2014). This threshold is reached when permeability exceeds approximately 10 -12 m 2 (Huettel and Rusch, 2000). In fine-grained sediments with permeabilities below this threshold, solute transport is dominated by molecular diffusion due to existing concentration gradients of solutes and the relatively low hydraulic conductivity of the sediments (Huettel et al., 2003). Diffusion in sediment porewater is a very slow process because the diffusing molecules must follow a tortuous path around the sediment grains (Huettel and Webster, 2001). In coarse-grained, permeable sediments with permeabilities above the threshold, advective processes dominate over diffusion.
Unidirectional and wave orbital water flows interacting with microscale topography (e.g. ripples and biogenic mounds) at the water–sediment interface lead to increased fluid exchange rates compared to exchange by molecular diffusion (Huettel et al., 1996; Precht and Huettel, 2003). Interaction of flows with surface microtopography increases oxygen penetration depths (Huettel and Rusch, 2000). The advective supply of oxygen to the sedimentary microbial community facilitates the effective remineralization of OC filtered into the surface layers of permeable sediment (Huettel et al., 2014). Because of advective porewater flows, permeable sediments may act as biocatalytic filters, notable for their high reaction rates, intense recycling, and extreme spatial and temporal dynamics of biogeochemical processes (Huettel et al., 2003; 2014).
In hydrodynamically quiet, depositional environments trawling may increase OC remineralisation, due to the oxygenation of sediments and redeposition of recently expulsed organic material back to the seabed. (Duplisea et al., 2001; Polymenakou et al., 2005; van de Velde et al., 2018). However, remineralization due to trawling activity may be limited in hydrodynamically active environments due to the removal of fauna and finer surface sediments, low OC due to resuspension and lateral/vertical transportation, and typically coarser sediments associated with deeper oxygen penetration and higher natural remineralisation rates. (Burdige, 2007; Huettel et al., 2014; Pusceddu et al., 2014; Tiano et al., 2019; De Borger et al., 2021; Morys et al., 2021). By comparing disturbance caused by trawling with natural disturbance levels due to hydrodynamics, it might be possible to identify areas where anthropogenic fishing disturbance lies beyond the bounds of natural variability (Diesing et al., 2013).
8.5 - Conclusion
Present estimations of the effect of trawling on OC remineralisation have many uncertainties, assumptions and simplifications. This is due to a lack of site-specific understanding of the complex interactions between local sediment (e.g. grain size and OC content and stability), environmental conditions (e.g. temperature and oxygenation), biology (e.g. production and bioturbation) and hydrology (e.g. sediment mixing and transport) that determine local OC contents and rates of remineralization. However, it may be postulated that hydrodynamically quiet, depositional environments with finer sediments, higher OC contents, and low levels of bioturbation, natural disturbance and oxidation may be expected to show higher levels of OC remineralisation in response to sediment disturbance from bottom trawling. Conversely, in hydrodynamically active environments with coarser sediments, greater oxygen penetration and naturally high OC remineralisation rates, trawling activity may be expected to have a smaller effect on carbon cycling. Presently undisturbed areas in deeper/up-welling or Arctic waters may be particularly sensitive, as increasing temperatures have also been linked to increased remineralization (Qi et al., 2020; Yamamoto-Kawai et al., 2009), depending on OC sediment concentration and if nutrients are limiting. These environments are also predicated to be some of the most sensitive to ocean acidification and so even small local change in seawater DIC and carbonate chemistry may have an impact on ecosystem function. More site specific in situ studies of carbon fluxes in response to trawling activity are needed in Norwegian waters, particularly when considering opening new areas to trawling.
8.6 - References
Adriano, S., Massimiliano, F., Sonia, C., Chiara, F., & Marcomini, A. (2005). Organic carbon changes in the surface sediments of the Venice lagoon. Environment International, 31(7), 1002–1010. https://doi.org/10.1016/j.envint.2005.05.010
Allen, J., & Clarke, K. (2007). Effects of demersal trawling on ecosystem functioning in the North Sea: A modelling study. Marine Ecology Progress Series, 336, 63–75. https://doi.org/10.3354/meps336063
Aller, R.C., Cochran, J.K. (2019). The critical role of Bioturbation for particle dynamics, priming potential, and organic C remineralization in marine sediments: Local and basin scales. Front. Earth Sci. 7, 157.
Amon, R.M.W., Benner, R. (1996). Bacterial utilization of different size classes of dissolved organic matter. Limnol. Oceanogr. 41, 41–51.
Atwood, T. B., Witt, A., Mayorga, J., Hammill, E., & Sala, E. (2020). Global patterns in marine sediment carbon stocks. Frontiers in Marine Science, 7, 165. https://doi.org/10.3389/fmars.2020.00165
Asper, V.L. (1987). Measuring the flux and sinking speed of marine snow aggregates, Deep Sea Research Part A. Oceanographic Research Papers, 34 (1), 1-17. https://doi.org/10.1016/0198-0149(87)90117-8.
Arndt, S., Jørgensen, B. B., LaRowe, D. E., Middelburg, J. J., Pancost, R. D., & Regnier, P. (2013). Quantifying the degradation of organic matter in marine sediments: A review and synthesis. Earth-Science Reviews, 123, 53–86. https://doi.org/10.1016/j.earscirev.2013.02.008
Bear J. (1972). Dynamics of Fluids in Porous Media. New York: Am. Elsevier
Bengtsson, M. M., Attermeyer, K., & Catalán, N. (2018). Interactive effects on organic matter processing from soils to the ocean: Are priming effects relevant in aquatic ecosystems? Hydrobiologia, 822(1), 1–17. https://doi.org/10.1007/s10750-018-3672-2
Berner, R. A. (1982). Burial of organic carbon and pyrite sulfur in the modern ocean; Its geochemical and environmental significance. American Journal of Science, 282(4), 451–473. https://doi.org/10.2475/ajs.282.4.451
Black, K.E., Smeaton, C., Turrell, W.R., Austin, W.E.N. (2022). Assessing the potential vulnerability of sedimentary carbon stores to bottom trawling disturbance within the UK EEZ. Front Mar Sci 9:892892. https://doi:10.3389/fmars.2022.892892
Bianchi, T. S. (2011). The role of terrestrially derived organic carbon in the coastal ocean: A changing paradigm and the priming effect. Proceedings of the National Academy of Sciences, 108(49), 19473–19481. https://doi.org/10.1073/pnas.1017982108
Bourgeois, S., Archambault, P., Witte, U. (2017). Organic matter remineralization in marine sediments: A pan-Arctic synthesis: Pan-Arctic Benthic Remineralization. Global Biogeochem. Cycles 31, 190–213.
Brin, L.D., Giblin, A.E. & Rich, J.J. (2015). Effects of experimental warming and carbon addition on nitrate reduction and respiration in coastal sediments. Biogeochemistry 125, 81–95. https://doi.org/10.1007/s10533-015-0113-4
Brodersen, K.E., Trevathan-Tackett, S.M., Nielsen, D.A., Connolly, R.M., Lovelock, C.E., Atwood, T.B., Macreadie, P.I. (2019). Oxygen consumption and sulfate reduction in Vegetated Coastal habitats: Effects of physical disturbance. Front. Mar. Sci. 6 (14).
Burdige, D. J. (2007). Preservation of organic matter in marine sediments: Controls, mechanisms, and an imbalance in sediment organic carbon budgets? Chemical Reviews, 107(2), 467–485. https://doi.org/10.1021/cr050347q
Capuzzo, E., Stephens, D., Silva, T., Barry, J., & Forster, R. M. (2015). Decrease in water clarity of the southern and central North Sea during the 20th century. Global Change Biology, 21(6), 2206–2214. https://doi.org/10.1111/gcb.12854
Chen, Z., Nie, T., Zhao, X., Li, J., Yang, B., Cui, D., Li, X. (2022). Organic carbon remineralization rate in global marine sediments: A review, Regional Studies in Marine Science, Volume 49, 102112, ISSN 2352-4855, https://doi.org/10.1016/j.rsma.2021.102112.
Cloern, J. E., Foster, S. Q., & Kleckner, A. E. (2014). Phytoplankton primary production in the world's estuarine-coastal ecosystems. Biogeosciences, 11(9), 2477–2501. https://doi.org/10.5194/bg112477-2014
Dauwe, B., Middelburg, J. J., & Herman, P. M. J. (2001). Effect of oxygen on the degradability of organic matter in subtidal and intertidal sediments of the North Sea area. Marine Ecology Progress Series, 215, 13–22. https://doi.org/10.3354/meps215013
De Borger, E., Tiano, J., Braeckman, U., Rijnsdorp, A. D., & Soetaert, K. (2021). Impact of bottom trawling on sediment biogeochemistry: A modelling approach. Biogeosciences, 18, 2539–2557. https://doi.org/10.5194/bg-2020-328
Deng, J., Zhu, W., Zhou, Y., Yin, Y. (2019). Soil organic carbon chemical functional groups under different revegetation types are coupled with changes in the microbial community composition and the functional genes. Forests 10 (240).
D’Hondt, S., Inagaki, F., Zarikian, C.A., Abrams, L.J., Dubois, N., Engelhardt, T., Evans, H., Ferdelman, T., Gribsholt, B., Harris, R.N., Hoppie, B.W., Hyun, J.-H., Kallmeyer, J., Kim, J., Lynch, J.E., McKinley, C.C., Mitsunobu, S., Morono, Y., Murray, R.W., Pockalny, R., Sauvage, J., Shimono, T., Shiraishi, F., Smith, D.C., Smith-Duque, C.E., Spivack, A.J., Steinsbu, B.O., Suzuki, Y., Szpak, M., Toffin, L., Uramoto, G., Yamaguchi, Y.T., Zhang, G., Zhang, X.-H., Ziebis, W. (2015). Presence of oxygen and aerobic communities from sea floor to basement in deep-sea sediments. Nat. Geosci. 8, 299–304.
Diesing, M., Kroger, S., Parker, R., Jenkins, C., Mason, C., & Weston, K. (2017). Predicting the standing stock of organic carbon in surface sediments of the North-West European continental shelf. Biogeochemistry, 135(1), 183–200. https://doi.org/10.1007/s1053 3-017-0310-4
Diesing M, Stephens D, Aldridge, J. (2013). A proposed method for assessing the extent of the seabed significantly affected by demersal fishing in the Greater North Sea. ICES J Mar Sci 70:1085–1096. https://doi.org/10.1093/icesjms/fst066
Diesing, M., Thorsnes, T., & Bjarnadóttir, L. R. (2021). Organic carbon densities and accumulation rates in surface sediments of the North Sea and Skagerrak. Biogeosciences, 18(6), 2139–2160. https://doi.org/10.5194/bg-18-2139-2021
Donis, D., McGinnis, D.F., Holtappels, M., Felden, J., Wenzhoefer, F. (2016). Assessing benthic oxygen fluxes in oligotrophic deep sea sediments (HAUSGARTEN observatory). Deep Sea Res. I: Oceanogr. Res. Pap. 111, 1–10.
Duplisea, D. E., Jennings, S., Malcolm, S. J., Parker, R., & Sivyer, D. B. (2001). Modelling potential impacts of bottom trawl fisheries on soft sediment biogeochemistry in the North Sea. Geochemical Transactions, 2(14), 112. https://doi.org/10.1186/1467-4866-2-112
Durrieu de Madron, X., Ferré, B., Le Corre, G., Grenz, C., Conan, P., PujoPay, M., Buscail, R., & Bodiot, O. (2005). Trawling-induced resuspension and dispersal of muddy sediments and dissolved elements in the Gulf of Lion (NW Mediterranean). Continental Shelf Research, 25(19–20), 2387–2409. https://doi.org/10.1016/j.csr.2005.08.002
Duursma, E.K. (1963). The production of dissolved organic matter in the sea, as related to the primary gross production of organic matter. Netherlands Journal of Sea Research, 2 (1), 85-94. https://doi.org/10.1016/0077-7579(63)90007-3
Ekdale, A., Bromley, R., & Pemberton, S. (1984). Effects of bioturbation on sediment properties. In Ichnology: The use of trace fossils in sedimentology and stratigraphy. Special Publications of SEPM: Society for Sedimentary Geology, USA.
Emerson, S. (2013). In: Sundquist, E.T., Broecker, W.S. (Eds.), Organic Carbon Preservation in Marine Sediments. In: Geophysical Monograph Series, American Geophysical Union, Washington, D.C.,
Nature 607, E1–E2. https://doi.org/10.1038/s41586-022-04775-1
Hinojosa, J.L., Moy, C.M., Stirling, C.H., Wilson, G.S., Eglinton, T.I. (2014). Carbon cycling and burial in New Zealand’s fjords. Geochem. Geophys. Geosyst. 15, 4047–4063.
Huettel, M. and Rusch, A. (2000). Transport and degradation of phytoplankton in permeable sediment, Limnol. Oceanogr., 45, 534–549, https://doi.org/10.4319/lo.2000.45.3.0534.
Huettel, M. and I. T. Webster. (2001). Porewater flow in permeable sediments, p. 144-179. In B. P. Boudreau and B. B. Jørgensen [eds.], The Benthic Boundary Layer. Oxford University Press.
Huettel, M., Berg, P., & Kostka, J. E. (2014). Benthic exchange and biogeochemical cycling in permeable sediments. Annual Review of Marine Science, 6(1), 23–51. https://doi.org/10.1146/annurev-marine-051413-012706
Huettel, M., Røy, H., Precht, E., and Ehrenhauss, S. (2003). Hydrodynamical impact on biogeochemical processes in aquatic sediments, Hydrobiologia, 494, 231–236.
Huettel, M., Ziebis,W., and Forster, S. (1996). Flow-induced uptake of particulate matter in permeable sediments, Limnol. Oceanogr., 41, 309–322, pp. 78–87.
Epstein, G., Middelburg, J. J., Hawkins, J. P., Norris, C. R., & Roberts, C. M. (2022). The impact of mobile demersal fishing on carbon storage in seabed sediments. Global Change Biology, 28, 2875–2894. https://doi.org/10.1111/gcb.16105
Epstein, G., Roberts, C.M. (2022). Identifying priority areas to manage mobile bottom fishing on seabed carbon in the UK. PLOS Clim 1:1–21. https://doi:10.1371/journal.pclm.0000059
Falcão, M., Gaspar, M. B., Caetano, M., Santos, M. N., & Vale, C. (2003). Short-term environmental impact of clam dredging in coastal waters (south of Portugal): Chemical disturbance and subsequent recovery of seabed. Marine Environmental Research, 56(5), 649–664. https://doi.org/10.1016/s0141-1136(03)00069-2
Fanning, K. A., Carder, K. L., & Betzer, P. R. (1982). Sediment resuspension by coastal waters: A potential mechanism for nutrient re-cycling on the ocean's margins. Deep Sea Research Part A. Oceanographic Research Papers, 29(8), 953–965. https://doi.org/10.1016/0198-0149(82)90020-6
Fiedler, B., Grundle, DS., Schutte, F., Karstensen, J., Loscher, C.R., Hauss, H., Wagner, H., Loginova, A., Kiko, R., Silva, P., Tauhua, T., Kortzinger, A. (2016). Oxygen utilization and downward carbon flux in an oxygen-depleted eddy in the eastern tropical North Atlantic. Biogeosciences, 13, 5633–5647, 2016 https://doi.org/10.5194/bg-13-5633-2016
Glud, R.N. (2008). Oxygen dynamics of marine sediments. Marine Biol. Res. 4, 243–289.
Graves C.A., Benson, L., Aldridge, J., Austin, W.E.N., Dal Molin. F., Fonseca, V.G., Hicks, N., Hynes, C., Kröger, S., Lamb, P.D., Mason, C., Powell, C., Smeaton, C., Wexler, S.K., Woulds, C. and Parker, R. (2022). Sedimentary carbon on the continental shelf: Emerging capabilities and research priorities for Blue Carbon. Front. Mar. Sci. 9:926215. https://doi:10.3389/fmars.2022.926215
Hilborn, R., Kaiser, M.J. (2022). A path forward for analysing the impacts of marine protected areas. https://doi.org/10.4319/lo.1996.41.2.0309.
Hulthe, G., Hulth, S., & Hall, P. O. J. (1998). Effect of oxygen on degradation rate of refractory and labile organic matter in continental margin sediments. Geochimica et Cosmochimica Acta, 8, 1319–1328. https://doi.org/10.1016/S0016-7037(98)00044-1
Jennings, S., Dinmore, T. A., Duplisea, D. E., Warr, K. J., & Lancaster, J. E. (2001). Trawling disturbance can modify benthic production processes. Journal of Animal Ecology, 70(3), 459–475. https://doi.org/10.1046/j.1365-2656.2001.00504.x
Jennings, S., Nicholson, M. D., Dinmore, T. A., & Lancaster, J. E. (2002). Effects of chronic trawling disturbance on the production of infaunal communities. Marine Ecology Progress Series, 243, 251–260. https://doi.org/10.3354/meps243251
Jessen, G.L., Lichtschlag, A., Ramette, A., Pantoja, S., Rossel, P.E., Schubert, C.J., Struck, U., Boetius, A. (2017). Hypoxia causes preservation of labile organic matter and changes seafloor microbial community composition (Black Sea). Sci. Adv. 3, e1601897.
Keil, R. (2017). Anthropogenic forcing of carbonate and organic carbon preservation in marine sediments. Annual Review of Marine Science, 9(1), 151–172. https://doi.org/10.1146/annurev-marine-010816-060724
Kaiser, M. J., Clarke, K. R., Hinz, H., Austen, M. C. V., Somerfield, P. J., & Karakassis, I. (2006). Global analysis of response and recovery of benthic biota to fishing. Marine Ecology Progress Series, 311, 1–14. https://doi.org/10.3354/meps311001
Kaiser, M. J., Collie, J. S., Hall, S. J., Jennings, S., & Poiner, I. R. (2002). Modification of marine habitats by trawling activities prognosis and solutions. Fish and Fisheries, 3, 114–136. https://doi.org/10.1046/j.1467-2979.2002.00079.x
Kleber, M., Lehmann, J. (2019). Humic substances extracted by alkali are invalid proxies for the dynamics and functions of organic matter in terrestrial and aquatic ecosystems. J. Environ. Qual. 48, 207–216.
Kristensen, E., Ahmed, S. I., & Devol, A. H. (1995). Aerobic and anaerobic decomposition of organic matter in marine sediment: Which is fastest? Limnology and Oceanography, 40(8), 1430–1437. https://doi.org/10.4319/lo.19126.96.36.1990
Kurian, S., Kessarkar, P.M., Rao, V.Purnachandra., Reshma, K., Sarkar, A., Pattan, J.N., Naqvi, S.W.A. (2020). Controls on organic matter distribution in oxygen minimum zone sediments from the continental slope off western India. J. Mar. Syst. 207, 103118.
LaRowe, D. E., Arndt, S., Bradley, J. A., Estes, E. R., Hoarfrost, A., Lang, S. Q., Lloyd, K. G., Mahmoudi, N., Orsi, W. D., Shah Walter, S. R., Steen, A. D., & Zhao, R. (2020). The fate of organic carbon in marine sediments—New insights from recent data and analysis. Earth Science Reviews, 204, 103146. https://doi.org/10.1016/j.earscirev. 2020.103146
Lee, T. R., Wood, W. T., & Phrampus, B. J. (2019). A machine learning (kNN) approach to predicting global seafloor total organic carbon. Global Biogeochemical Cycles, 33(1), 37–46. https://doi.org/10.1029/2018gb005992
Legge, O., Johnson, M., Hicks, N., Jickells, T., Diesing, M., Aldridge, J., Andrews, J., Artioli, Y., Bakker, D. C. E., Burrows, M. T., Carr, N., Cripps, G., Felgate, S. L., Fernand, L., Greenwood, N., Hartman, S., Kröger, S., Lessin, G., Mahaffey, C., … Williamson, P. (2020). Carbon on the Northwest European Shelf: Contemporary budget and future influences. Frontiers in Marine Science, 7, Article 143. https://doi.org/10.3389/fmars.2020.00143
Luisetti, T., Turner, R. K., Andrews, J. E., Jickells, T. D., Kröger, S., Diesing, M., Paltriguera, L., Johnson, M. T., Parker, E. R., Bakker, D. C. E., & Weston, K. (2019). Quantifying and valuing carbon flows and stores in coastal and shelf ecosystems in the UK. Ecosystem Services, 35, 67–76. https://doi.org/10.1016/j.ecoser.2018.10.013
Malinverno, A., Martinez, E.A. (2015). The effect of temperature on organic carbon degradation in marine sediments. Sci. Rep. 5 (17861).
Martín, J., Puig, P., Masque, P., Palanques, A., & Sanchez-Gomez, A. (2014a). Impact of bottom trawling on deep-sea sediment properties along the flanks of a submarine canyon. PLoS One, 9(8), e104536. https://doi.org/10.1371/journal.pone.0104536
Martín, J., Puig, P., Palanques, A., & Giamportone, A. (2014b). Commercial bottom trawling as a driver of sediment dynamics and deep seascape evolution in the Anthropocene. Anthropocene, 7, 1–15. https://doi.org/10.1016/j.ancene.2015.01.002
Mengual, B., Cayocca, F., Le Hir, P., Draye, R., Laffargue, P., Vincent, B., & Garlan, T. (2016). Influence of bottom trawling on sediment resuspension in the ‘Grande-Vasière’ area (Bay of Biscay, France). Ocean Dynamics, 66(9), 1181–1207. https://doi.org/10.1007/s10236-016-0974-7
Meysman, F. J. R., Middelburg, J. J., & Heip, C. H. R. (2006). Bioturbation: A fresh look at Darwin's last idea. Trends in Ecology & Evolution, 21(12), 688–695. https://doi.org/10.1016/j.tree.2006.08.002
Middelburg, J. J. (2018). Reviews and syntheses: To the bottom of carbon processing at the seafloor. Biogeosciences, 15(2), 413–427. https://doi.org/10.5194/bg-15-413-2018
Morys, C., Brüchert, V., & Bradshaw, C. (2021). Impacts of bottom trawling on benthic biogeochemistry: An experimental field study. Marine Environmental Research, 169, 105384. https://doi.org/10.1016/j.marenvres.2021.105384
Nedwell, D.B. (1999). Effect of low temperature on microbial growth: lowered affinity for substrates limits growth at low temperature. FEMS Microbiol. Ecol. 30, 101–111.
Nierop, K.G.J., Reichart, G.-J., Veld, H., Sinninghe Damsté, J.S. (2017). The influence of oxygen exposure time on the composition of macromolecular organic matter as revealed by surface sediments on the Murray Ridge (Arabian Sea). Geochim. Cosmochim. Acta 206, 40–56.
Oberle, F. K. J., Storlazzi, C. D., & Hanebuth, T. J. J. (2016a). What a drag: Quantifying the global impact of chronic bottom trawling on continental shelf sediment. Journal of Marine Systems, 159, 109–119. https://doi.org/10.1016/j.jmarsys.2015.12.007
Oberle, F. K. J., Swarzenski, P. W., Reddy, C. M., Nelson, R. K., Baasch, B., & Hanebuth, T. J. J. (2016b). Deciphering the lithological consequences of bottom trawling to sedimentary habitats on the shelf. Journal of Marine Systems, 159, 120–131. https://doi.org/10.1016/j. jmarsys.2015.12.008
Olsgard, F., Schaanning, M.T., Widdicombe, S., Kendall, M.A., Austen, M.C. (2008). Effects of bottom trawling on ecosystem functioning, Journal of Experimental Marine Biology and Ecology, 366, (1–2) 123-133, https://doi.org/10.1016/j.jembe.2008.07.036.
Palanques, A., Puig, P., Guillén, J., Demestre, M., & Martín, J. (2014). Effects of bottom trawling on the Ebro continental shelf sedimentary system (NW Mediterranean). Continental Shelf Research, 72, 83–98. https://doi.org/10.1016/j.csr.2013.10.008
Palanques, A., Guillén, J., & Puig, P. (2001). Impact of bottom trawling on water turbidity and muddy sediment of an unfished continental shelf. Limnology and Oceanography, 46(5), 1100–1110. https://doi.org/10.4319/lo.2001.46.5.1100
Paradis, S., Goñi, M., Masqué, P., Durán, R., Arjona-Camas, M., Palanques, A., & Puig, P. (2021). Persistence of biogeochemical alterations of deep-sea sediments by bottom trawling. Geophysical Research Letters, 48(2), e2020GL091279. https://doi.org/10.1029/2020g l091279
Polymenakou, P. N., Pusceddu, A., Tselepides, A., Polychronaki, T., Giannakourou, A., Fiordelmondo, C., Hatziyanni, E., & Danovaro, R. (2005). Benthic microbial abundance and activities in an intensively trawled ecosystem (Thermaikos Gulf, Aegean Sea). Continental Shelf Research, 25(19–20), 2570–2584. https://doi.org/10.1016/j.csr.2005.08.018
Precht, E. and Huettel, M. (2003). Advective pore-water exchange driven by surface gravity waves and its ecological implications, Limnol. Oceanogr., 48, 1674–1684.
Pusceddu, A., Bianchelli, S., & Danovaro, R. (2015). Quantity and biochemical composition of particulate organic matter in a highly trawled area (Thermaikos Gulf, Eastern Mediterranean Sea). Advances in Oceanography and Limnology, 6(1/2), 21–34. https://doi.org/10.4081/aiol.2015.5448
Pusceddu, A., Bianchelli, S., Martin, J., Puig, P., Palanques, A., Masque, P., & Danovaro, R. (2014). Chronic and intensive bottom trawling impairs deep-sea biodiversity and ecosystem functioning. Proceedings of the National Academy of Sciences of the United States of America, 111(24), 8861–8866. https://doi.org/10.1073/pnas.1405454111
Qi, D., Chen, B., Chen, L., Lin, H., Gao, Z., Sun, H., Zhang, Y., Sun, X., Cai, W. (2020). Coastal acidification induced by biogeochemical processes driven by sea-ice melt in the western Arctic ocean. Polar Sci. 23, 100504.
Queirós, A. M., Hiddink, J. G., Kaiser, M. J., & Hinz, H. (2006). Effects of chronic bottom trawling disturbance on benthic biomass, production and size spectra in different habitats. Journal of Experimental Marine Biology and Ecology, 335(1), 91–103. https://doi.org/10.1016/j.jembe.2006.03.001
Ramirez-Llodra, E., Rinde, E., Gundersen, H. Fagerli CW, Fredriksen S, Gitmark JK, Norling K, Walday MG, Norderhaug KM. (2016). A snap shot of the short-term response of crustaceans to macrophyte detritus in the deep Oslofjord. Sci Rep 6, 23800. https://doi.org/10.1038/srep23800
Riekenberg, P. M., Oakes, J. M., & Eyre, B. D. (2020). Shining light on priming in euphotic sediments: Nutrient enrichment stimulates export of stored organic matter. Environmental Science & Technology, 54(18), 11165–11172. https://doi.org/10.1021/acs.est.0c01914
Roberts, C. M. (2007). The unnatural history of the sea. Island Press.
Roussel, E.G., Cragg, B.A., Webster, G., Sass, H., Tang, X., Williams, A.S., Gorra, R., Weightman, A.J., Parkes, R.J. (2015). Complex coupled metabolic and prokaryotic community responses to increasing temperatures in anaerobic marine sediments: critical temperatures and substrate changes. FEMS Microbiol.Ecol. 91, fiv084.
Ruffin, K. K. (1998). The persistence of anthropogenic turbidity plumes in a shallow water estuary. Estuarine, Coastal and Shelf Science, 47(5), 579–592. https://doi.org/10.1006/ecss.1998.0366
Rühl, S., Thompson, C., Queirós, A. M., & Widdicombe, S. (2020). Missing links in the study of solute and particle exchange between the sea floor and water column. ICES Journal of Marine Science, 77(5), 1602– 1616. https://doi.org/10.1093/icesjms/fsaa060
Sala, E., Mayorga, J., Bradley, D., Cabral, R. B., Atwood, T. B., Auber, A., Cheung, W., Costello, C., Ferretti, F., Friedlander, A. M., Gaines, S. D., Garilao, C., Goodell, W., Halpern, B. S., Hinson, A., Kaschner, K., Kesner-Reyes, K., Leprieur, F., McGowan, J., Lubchenco, J. (2021). Protecting the global ocean for biodiversity, food and climate. Nature, 592, 397–402. https://doi.org/10.1038/s41586-021-03371-z
Sanches, L. F., Guenet, B., dos Anjos Cristiano Marino, N., and de Assis Esteves, F. (2021). Exploring the drivers controlling the priming effect and its magnitude in aquatic systems. J. Geophys.ical Res.: Biogeosci. 126, e2020JG006201. https://doi:10.1029/2020JG006201
Sciberras, M., Hiddink, J. G., Jennings, S., Szostek, C. L., Hughes, K. M., Kneafsey, B., Clarke, L. J., Ellis, N., Rijnsdorp, A. D., McConnaughey, R. A., Hilborn, R., Collie, J. S., Pitcher, C. R., Amoroso, R. O., Parma, A. M., Suuronen, P., & Kaiser, M. J. (2018). Response of benthic fauna to experimental bottom fishing: A global meta-analysis. Fish and Fisheries, 19(4), 698–715. https://doi.org/10.1111/faf.12283
Seiter, K., Hensen, C., Schröter, J., & Zabel, M. (2004). Organic carbon content in surface sediments—Defining regional provinces. Deep Sea Research Part I: Oceanographic Research Papers, 51(12), 2001– 2026. https://doi.org/10.1016/j.dsr.2004.06.014
Seiter, K., Hensen, C., Zabel, M. (2005). Benthic carbon mineralization on a global scale. Global Biogeochem. Cycles 19, GB1010.
Smeaton, C., Hunt, C. A., Turrell, W. R., & Austin, W. E. N. (2021). Marine sedimentary carbon stocks of the United Kingdom’s exclusive economic zone. Frontiers in Earth Science, 9, 50. https://doi.org/10.3389/feart.2021.593324
Snelgrove, P. V. R., Soetaert, K., Solan, M., Thrush, S., Wei, C.-L., Danovaro, R., Fulweiler, R. W., Kitazato, H., Ingole, B., Norkko, A., Parkes, R. J., & Volkenborn, N. (2018). Global carbon cycling on a heterogeneous seafloor. Trends in Ecology & Evolution, 33(2), 96– 105. https://doi.org/10.1016/j.tree.2017.11.004
Stumm-Zollinger, E. (1968). Substrate utilization in heterotrophic bacterial communities. J. Water Pollution Ctrl. Fed., 40, 213-229.
Tabuchi, K., Kojima, H. & Fukui, M. (2010). Seasonal Changes in Organic Matter Mineralization in a Sublittoral Sediment and Temperature-Driven Decoupling of Key Processes. Microb Ecol 60, 551–560. https://doi.org/10.1007/s00248-010-9659-9
Thrush, S. F., & Dayton, P. K. (2002). Disturbance to marine benthic habitats by trawling and dredging: Implications for marine biodiversity. Annual Review of Ecology and Systematics, 33(1), 449–473. https://doi.org/10.1146/annurev.ecolsys.33.010802.150515
Tiano, J. C., van der Reijden, K. J., O'Flynn, S., Beauchard, O., van der Ree, S., van der Wees, J., Ysebaert, T., & Soetaert, K. (2020). Experimental bottom trawling finds resilience in large-bodied infauna but vulnerability for epifauna and juveniles in the Frisian Front. Marine Environmental Research, 159, 104964. https://doi.org/10.1016/j.marenvres.2020.104964
Tiano, J. C., Witbaard, R., Bergman, M. J. N., van Rijswijk, P., Tramper, A., van Oevelen, D., & Degraer, S. (2019). Acute impacts of bottom trawl gears on benthic metabolism and nutrient cycling. ICES Journal of Marine Science, 76(6), 1917–1930. https://doi.org/10.1093/icesjms/fsz060
Tietjen, T., G. Wetzel, R. (2003). Extracellular enzyme-clay mineral complexes: Enzyme adsorption, alteration of enzyme activity, and protection from photodegradation. Aquat. Ecol. 37, 331–339.
Tillin, H. M., Hiddink, J. G., Jennings, S., & Kaiser, M. J. (2006). Chronic bottom trawling alters the functional composition of benthic invertebrate communities on a sea-basin scale. Marine Ecology Progress Series, 318, 31–45. https://doi.org/10.3354/meps318031
Trevathan-Tackett, S.M., Thomson, A.C.G., Ralph, P.J., Macreadie, P.I. (2018). Fresh carbon inputs to seagrass sediments induce variable microbial priming responses. Sci. Total Environ. 621, 663–669.
Trimmer, M., Petersen, J., Sivyer, D. B., Mills, C., Young, E., & Parker, E. R. (2005). Impact of long-term benthic trawl disturbance on sediment sorting and biogeochemistry in the southern North Sea. Marine Ecology Progress Series, 298, 79–94. https://doi.org/10.3354/meps2 98079
Turner, J. T. (2015). Zooplankton fecal pellets, marine snow, phytodetritus and the ocean's biological pump. Progress in Oceanography, 130, 205–248. https://doi.org/10.1016/j.pocean.2014.08.005
Van der Molen, J., Aldridge, J. N., Coughlan, C., Parker, E. R., Stephens, D., & Ruardij, P. (2012). Modelling marine ecosystem response to climate change and trawling in the North Sea. Biogeochemistry, 113(1–3), 213–236. https://doi.org/10.1007/s10533-012-9763-7
Van de Velde, S., Van Lancker, V., Hidalgo-Martinez, S., Berelson, W. M., & Meysman, F. J. R. (2018). Anthropogenic disturbance keeps the coastal seafloor biogeochemistry in a transient state. Scientific Reports, 8(1), 5582. https://doi.org/10.1038/s41598-018-23925-y
Van Kaam-Peters, H.M.E., Schouten, S., Köster, J., Sinninghe Damstè, J.S. (1998). Controls on the molecular and carbon isotopic composition of organic matter deposited in a Kimmeridgian euxinic shelf sea: evidence for preservation of carbohydrates through sulfurisation. Geochim. Cosmochim. Acta., 62, 3259–3283.
Van Nugteren, P., Moodley, L., Brummer, G.-J., Heip, C. H. R., Herman, P. M. J., & Middelburg, J. J. (2009). Seafloor ecosystem functioning: The importance of organic matter priming. Marine Biology, 156(11), 2277–2287. https://doi.org/10.1007/s00227-009-1255-5
Weston, N.B., Joye, S.B. (2005). Temperature-driven decoupling of key phases of organic matter degradation in marine sediments. Proc. Natl. Acad. Sci. 102, 17036–17040.
Wilkinson, G. M., Besterman, A., Buelo, C., Gephart, J., & Pace, M. L. (2018). A synthesis of modern organic carbon accumulation rates in coastal and aquatic inland ecosystems. Scientific Reports, 8(1), 15736. https://doi.org/10.1038/s41598-018-34126-y
Wilson AM, Huettel M, Klein S. (2008). Grain size and depositional environment as predictors of permeability in coastal marine sands. Estuar Coast Shelf Sci 80:193–199. https://doi.org/10.1016/j.ecss.2008.06.011
Xue, J., Cai, W., Hu, X., Huang, W., Lohrenz, S.E., Gundersen, K. (2015). Temporal variation and stoichiometric ratios of organic matter remineralization in bottom waters of the northern Gulf of Mexico during late spring and summer. J. Geophys. Res. Oceans 120, 8304–8326.
Yamamoto-Kawai, M., McLaughlin, F.A., Carmack, E.C., Nishino, S., Shimada, K. (2009). Aragonite undersaturation in the Arctic ocean: effects of ocean acidification and sea ice melt. Science 326, 1098–1100.
Zang, H., Blagodatskaya, E., Wen, Y., Shi, L., Cheng, F., Chen, H., Zhao, B., Zhang, F., Fan, M., Kuzyakov, Y. (2020). Temperature sensitivity of soil organic matter mineralization decreases with long-term N fertilization: Evidence from four Q10 estimation approaches. Land Degrad. Dev. 31, 683–693.
Zhao, B., Yao, P., Bianchi, T.S., Arellano, A.R., Wang, X., Yang, J., Su, R., Wang, J., Xu, Y., Huang, X., Chen, L., Ye, J., Yu, Z. (2018). The remineralization of sedimentary organic carbon in different sedimentary regimes of the Yellow and East China Seas. Chemical Geology, 495, 104-117. https://doi.org/10.1016/j.chemgeo.2018.08.012.
Zimmerman, A.R., Chorover, J., Goyne, K.W., Brantley, S.L. (2004). Protection of mesopore-adsorbed organic matter from enzymatic degradation. Environ. Sci. Technol. 38, 4542–4548.