7 - Litteratur
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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.
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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.
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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
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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.
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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.
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