Ocean acidification reduces thallus strength in a non-calcifying foundation seaweed.

Climate change is causing unprecedented changes in terrestrial and aquatic ecosystems through the emission of greenhouse gases, including carbon dioxide (CO2). Approximately 30% of CO2 is taken up by the ocean ('ocean acidification', OA)1, which has profound effects on foundation seaweed species. Negative physical effects on calcifying algae are clear2, but studies on habitat-forming fleshy seaweeds have mainly focused on growth and less on thallus strength3,4. We exposed the habitat-forming brown seaweed Fucus vesiculosus to OA corresponding to projected climate change effects for the year 2100, and observed reduced apical thallus strength and greater loss of exposed individuals in the field. The tissue contained less calcium and magnesium, both of which are important for creating structural alginate matrices. Scanning electron microscopy (SEM) revealed tissue voids in the OA samples that were not present in seaweeds grown under ambient pCO2. We conclude that under OA, weakened F. vesiculosus will be at a significantly higher risk of physical damage and detachment.

Are silicone foul-release coatings a viable and environmentally sustainable alternative to biocidal antifouling coatings in the Baltic Sea region?

To combat unwanted fouling on immersed hulls, biocidal antifouling coatings are commonly applied to vessels trafficking the Baltic Sea. Here, the efficacy, environmental sustainability and market barriers of silicone foul-release coatings (FRCs) was assessed for this region to evaluate their viability as replacements for biocidal coatings. Coated panels were exposed statically over a 1 year period at three locations in the Baltic Sea region to assess the long-term performance of a biocide-free FRC and two copper coatings. The FRC was found to perform equally well or significantly better than the copper coatings. Even though most silicone FRCs on the market are biocide-free, a review of the literature regarding toxic effects and the identity and environmental fate of leachables shows that they may not be completely environmentally benign, simply for the lack of biocides. Nonetheless, FRCs are substantially less toxic compared to biocidal antifouling coatings and their use should be promoted.

Effect of ship hull form on the resistance penalty from biofouling.

Hull biofouling is a well-known problem for the shipping industry, leading to increased resistance and fuel consumption. Considering that the effects of hull form on resistance are known to be higher for a less slender hull, it is hypothesised in this paper that the effect of biofouling roughness on resistance is also dependent on the hull form. To test this hypothesis, previously reported full-scale numerical results on a containership are re-analysed. Form effects on roughness penalties, corresponding to KΔCT = 0.058 ± 0.025, are observed at a low speed (19 knots, Res = 2.29 × 109), which are however cancelled out by traditionally neglected roughness effects on wave-making resistance at a higher speed (24 knots, Res = 2.89 × 109). It is concluded that hull form effects on biofouling penalties can be significant at low speeds, though not generalisable for higher speeds, namely when wave-making resistance corresponds to ≥ 29% of total resistance.

Adaptation to dislodgement risk on wave-swept rocky shores in the snail Littorina saxatilis.

The periwinkle Littorina saxatilis has repeatedly evolved both a small, fragile and globose "wave ecotype" confined to wave-swept shores and a large, robust and elongated "crab ecotype" found in nearby crab-rich but less-exposed shores. This phenotypic divergence is assumed to reflect, in some part, local adaptation to wave exposure, but this hypothesis has received incomplete experimental testing. Here, we report a test of the prediction that the wave ecotype has a higher capacity to resist water flow than the crab ecotype. We sampled snails along a crab-wave transect and measured their resistance to dislodgement in a high-speed water flume with water speeds that match those of breaking waves in a range of relevant field conditions. Snails from the wave environment were consistently more resistant to water flow than snails from the crab environment and high resistance was positively correlated with the surface area of the foot and the area of the outer aperture contour both relative to shell size, and to the extent of lateral shell compression. In a separate experiment, we found that snails raised in still water in a common garden showed higher resistance to water flow if originating from a wave environment than from a crab environment, and this was true both at juvenile (2 weeks) and adult (10 months) developmental stages. This result suggests genetic control of a distinct "wave adapted" phenotype, likely to be maintained under strong divergent selection between the two adjacent habitats.

Instantaneous Flow Structures and Opportunities for Larval Settlement: Barnacle Larvae Swim to Settle.

Water flow affects settlement of marine larvae on several scales. At the smallest scale local flow regime may control the probability of adhesion to the substrate. Our aim was to mechanistically understand the transition from suspended to attached larvae in turbulent flow. Recently it was proposed that opportunities for larval settlement in turbulent boundary layers depend on time windows with suitable instantaneous flow properties. In flume flow we characterized the proportion of suitable time windows in a series of flow velocities with focus on the near-bed flow. The change in the proportion of potential settling windows with increasing free-stream velocities was compared to the proportion of temporary attachment of barnacle cypris larvae at different flow velocities. We found large instantaneous flow variations in the near-bed flow where cyprid attachment took place. The probability of temporary attachment in cyprids declined with local flow speed and this response was compatible with a settling window lasting at least 0.1 s with a maximum local flow speed of 1.9-2.4 cm s-1. Cyprids swam against the near-bed flow (negative rheotaxis) and the swimming speed (1.8 cm s-1) was close to the critical speed that permitted temporary attachment. We conclude that temporary attachment in barnacle cyprids requires upstream swimming to maintain a fixed position relative to the substrate for at least 0.1 s. This behaviour may explain the ability of barnacles to recruit to high-flow environments and give cyprids flexibility in the pre-settlement choice of substrates based on flow regime.

Discovery of symbiotic nitrogen fixation and chemoautotrophy in cold-water corals.

Cold-water corals (CWC) are widely distributed around the world forming extensive reefs at par with tropical coral reefs. They are hotspots of biodiversity and organic matter processing in the world's deep oceans. Living in the dark they lack photosynthetic symbionts and are therefore considered to depend entirely on the limited flux of organic resources from the surface ocean. While symbiotic relations in tropical corals are known to be key to their survival in oligotrophic conditions, the full metabolic capacity of CWC has yet to be revealed. Here we report isotope tracer evidence for efficient nitrogen recycling, including nitrogen assimilation, regeneration, nitrification and denitrification. Moreover, we also discovered chemoautotrophy and nitrogen fixation in CWC and transfer of fixed nitrogen and inorganic carbon into bulk coral tissue and tissue compounds (fatty acids and amino acids). This unrecognized yet versatile metabolic machinery of CWC conserves precious limiting resources and provides access to new nitrogen and organic carbon resources that may be essential for CWC to survive in the resource-depleted dark ocean.

Natural populations of shipworm larvae are attracted to wood by waterborne chemical cues.

The life cycle of many sessile marine invertebrates includes a dispersive planktonic larval stage whose ability to find a suitable habitat in which to settle and transform into benthic adults is crucial to maximize fitness. To facilitate this process, invertebrate larvae commonly respond to habitat-related chemical cues to guide the search for an appropriate environment. Furthermore, small-scale hydrodynamic conditions affect dispersal of chemical cues, as well as swimming behavior of invertebrate larvae and encounter with potential habitats. Shipworms within the family Teredinidae are dependent on terrestrially derived wood in order to complete their life cycle, but very little is known about the cues and processes that promote settlement. We investigated the potential for remote detection of settling substrate via waterborne chemical cues in teredinid larvae through a combination of empirical field and laboratory flume experiments. Natural populations of teredinid larvae were significantly more abundant close to wooden structures enclosed in plankton net compared to empty control nets, clearly showing that shipworm larvae can sense and respond to chemical cues associated with suitable settling substrate in the field. However, the flume experiments, using ecologically relevant flow velocities, showed that the boundary layer around experimental wooden panels was thin and that the mean flow velocity exceeded larval swimming velocity approximately 5 mm (≈ 25 larval body lengths) from the panel surface. Therefore, we conclude that the scope for remote detection of waterborne cues is limited and that the likely explanation for the higher abundance of shipworm larvae associated with the wooden panels in the field is a response to a cue during or after attachment on, or very near, the substrate. Waterborne cues probably guide the larva in its decision to remain attached and settle, or to detach and continue swimming and drifting until the next encounter with a solid substrate.

Embryogenesis and larval biology of the cold-water coral Lophelia pertusa.

Cold-water coral reefs form spectacular and highly diverse ecosystems in the deep sea but little is known about reproduction, and virtually nothing about the larval biology in these corals. This study is based on data from two locations of the North East Atlantic and documents the first observations of embryogenesis and larval development in Lophelia pertusa, the most common framework-building cold-water scleractinian. Embryos developed in a more or less organized radial cleavage pattern from ∼ 160 µm large neutral or negatively buoyant eggs, to 120-270 µm long ciliated planulae. Embryogenesis was slow with cleavage occurring at intervals of 6-8 hours up to the 64-cell stage. Genetically characterized larvae were sexually derived, with maternal and paternal alleles present. Larvae were active swimmers (0.5 mm s(-1)) initially residing in the upper part of the water column, with bottom probing behavior starting 3-5 weeks after fertilization. Nematocysts had developed by day 30, coinciding with peak bottom-probing behavior, and possibly an indication that larvae are fully competent to settle at this time. Planulae survived for eight weeks under laboratory conditions, and preliminary results indicate that these planulae are planktotrophic. The late onset of competency and larval longevity suggests a high dispersal potential. Understanding larval biology and behavior is of paramount importance for biophysical modeling of larval dispersal, which forms the basis for predictions of connectivity among populations.

Resistance of Lophelia pertusa to coverage by sediment and petroleum drill cuttings.

In laboratory experiments, the cold-water coral Lophelia pertusa was exposed to settling particles. The effects of reef sediment, petroleum drill cuttings and a mix of both, on the development of anoxia at the coral surface were studied using O2, pH and H2S microsensors and by assessing coral polyp mortality. Due to the branching morphology of L. pertusa and the release of coral mucus, accumulation rates of settling material on coral branches were low. Microsensors detected H2S production in only a few samples, and sulfate reduction rates of natural reef sediment slurries were low (<0.3 nmol S cm(-3) d(-1)). While the exposure to sediment clearly reduced the coral's accessibility to oxygen, L. pertusa tolerated both partial low-oxygen and anoxic conditions without any visible detrimental short-term effect, such as tissue damage or death. However, complete burial of coral branches for >24 h in reef sediment resulted in suffocation.

Tolerance to long-term exposure of suspended benthic sediments and drill cuttings in the cold-water coral Lophelia pertusa.

The cold-water coral Lophelia pertusa was exposed to suspended particles (<63 µm) for 12 weeks. Skeletal growth was significantly lower under exposure concentrations of ∼25 mg l⁻¹ than ∼5 mg l⁻¹ and there was a trend of lower growth rates when exposed to water-based drill cuttings than to natural benthic sediment. Polyp extension was less in corals exposed to higher material concentrations, which provides a possible explanation for observed skeletal growth differences between particle concentrations. Particle exposure had no significant impact on respiration or proportions of tissue and fatty acids in corals. The volume of additional cleaning mucus released by exposed corals was low and release did not significantly affect coral energy expenditure. Our results indicate that L. pertusa polyps can deal comparatively well with enhanced particle deposition rates and suspended matter concentrations. However, a small pilot experiment indicated that coral larvae might be particularly vulnerable to high particle concentrations.

Sedimentation on the cold-water coral Lophelia pertusa: cleaning efficiency from natural sediments and drill cuttings.

Anthropogenic threats to cold-water coral reefs are trawling and hydrocarbon drilling, with both activities causing increased levels of suspended particles. The efficiency of Lophelia pertusa in rejecting local sediments and drill cuttings from the coral surface was evaluated and found not to differ between sediment types. Further results showed that the coral efficiently removed deposited material even after repeated exposures, indicating an efficient cleaning mechanism. In an experiment focusing on burial, fine-fraction drill cuttings were deposited on corals over time. Drill cutting covered coral area increased with repeated depositions, with accumulation mainly occurring on and adjacent to regions of the coral skeleton lacking tissue cover. Tissue was smothered and polyp mortality occurred where polyps became wholly covered by material. Burial of coral by drill cuttings to the current threshold level used in environmental risk assessment models by the offshore industry (6.3mm) may result in damage to L. pertusa colonies.

Barnacle larvae actively select flow environments supporting post-settlement growth and survival.

Many marine dispersive propagules select specific settlement sites based on a range of environmental cues. However, the link between larval choice and post-settlement growth and survival is still poorly understood. Here we show that cypris larvae of the barnacle Balanus improvisus actively reject surfaces exposed to local flow speeds exceeding 5-10 cm/s. Field experiments show that post-settlement growth and survival decline in freestream flows above 15 cm/s. Moreover, studies in flume flow at local speeds exceeding 10 cm/s reveal that early juveniles show reduced feeding rates caused by deformation of the cirral fan, reduced retention efficiency, and a decrease in time spent feeding. We conclude that cypris larvae actively reject flow environments that will be suboptimal for suspension feeding in the early post-settlement phase. Our study suggests that larval choice can be adaptively connected to a specific part of the life cycle, in this case the very sensitive time after metamorphosis.