From sink to source: Dynamic of greenhouse gases emissions from beach wrack accumulations in a temperate coastal bay.

Coastal ecosystems such as salt marshes, seagrass meadows, and kelp forests contribute to climate regulation as carbon sinks. However, coastal ecosystems may act as carbon sources as beach wrack accumulations may release greenhouse gases (GHG) during decomposition. The magnitude of GHG emissions of beach wrack accumulations under natural conditions are poorly understood, hampering accurate blue carbon accountings. In this study, we assessed the spatio-temporal variability and environmental factors driving CO2, CH4 and N2O emissions from beach wrack accumulations on a temperate sandy beach. Beach wrack accumulations, dominated by Zostera marina and opportunistic brown macroalgae, presented variable spatio-temporal dynamics. Annual beach wrack GHG emissions achieved up to 77,915 mg m-2 d-1 CO2e (CO2 equivalents) and varied largely throughout the study period due to interactive effects of temperature, wave exposure, beach wrack biomass moisture, abundance, and species composition. Our findings showed that methane emissions in new, freshly deposited, and in drifting wrack in the water reached up to 100 mg m-2 d-1, representing up to 57 % of annual CO2e emissions occurring throughout the year. Nitrous oxide emissions were highly variable and comprised a minor extent (i.e., up to 4 %) of annual CO2e emissions. Together, wrack CH4 and N2O emissions provided 13.69 g CO2 m-2 per year to the atmosphere. Our findings indicate that excessive opportunistic macroalgae biomass driven by eutrophication may explain increased CO2 and N2O emissions. We conclude that whilst beach wrack depositions are a natural and essential part of coastal ecosystems, they may provide an extra source of GHG to the atmosphere, potentially counteracting the role of vegetated coastal ecosystems as carbon sinks.

Natural Algaecide Sphingosines Identified in Hybrid Straw Decomposition Driven by White-Rot Fungi.

Harmful algal blooms (HABs), which are promoted by eutrophication and intensified by global warming, occur worldwide. Allelochemicals, which are natural chemicals derived from plants or microbes, are emerging weapons to eliminate these blooms. However, the cost and technical challenges have limited the discovery of novel antialgal allelochemicals. Herein, the decomposition of agricultural straws is manipulated by white-rot fungi and achieved elevated antialgal efficiency. The transcriptomic analysis reveals that nutrient limitation activated fungal decomposition. By using a comparative nontarget metabolomics approach, a new type of allelochemical sphingosines (including sphinganine, phytosphingosine, sphingosine, and N-acetylsphingosine) is identified. These novel natural algaecides exhibit superior antialgal capability, with as high as an order of magnitude lower effective concentration on blooming species than other prevalent allelochemicals. The co-expression relationship between transcriptomic and metabolomic results indicate that sphinganine is strongly correlated with the differentially expressed lignocellulose degradation unigenes. The algal growth suppression is triggered by the activation of programmed cell death, malfunction of algal photosystem and antioxidant system, the disruption on CO2 assimilation and light absorption. The sphingosines reported here are a new category of allelochemicals in addition to the well-known antialgal natural chemicals, which are potential species-specific agents for HABs control identified by multi-omics methodology.

Effects of depth and overgrowth of ephemeral macroalgae on a remote subtidal NE Atlantic eelgrass (Zostera marina) community.

We conducted a short-term field sampling complemented with time integrating stable isotope analysis to holistically investigate status and ecological interactions in a remote NE Atlantic Zostera marina meadow. We found high nutrient water concentrations, large biomass of fast-growing, ephemeral macroalgae, low abundance, and biodiversity of epifauna and a food web with thornback ray (Raja clavata) as intermediate and cod (Gadus morhua) as top predator. We observed no variation with increasing depth (3.5-11 m) except for decreasing shoot density and biomass of Zostera and macroalgae. Our results indicate that the Finnøya Zostera ecosystem is eutrophicated. During the past three to four decades, nutrients from aquaculture have steadily increased to reach 75% of anthropogenic input while the coastal top predator cod has decreased by 50%. We conclude that bottom-up regulation is a predominant driver of change since top-down regulation is generally weak in low density and exposed Zostera ecosystems such as Finnøya.

New Insights into the Microplastic Enrichment in the Blue Carbon Ecosystem: Evidence from Seagrass Meadows and Mangrove Forests in Coastal South China Sea.

Microplastics were recently found to aggregate in the blue carbon ecosystems (BCEs), which are known for their ability to store carbon by slowing down the water flow. However, evidence is largely lacking on how the accumulation of microplastics is related to carbon sequestration in BCEs and if this trap effect is driven by its biological characteristics. In this study, the trap effect of microplastics by BCEs was evaluated for various seagrasses (Zostera japonica, Halophila ovalis, and Halophila beccarii) and mangroves (Aegiceras corniculatum and Avicennia marina). Significant accumulation was found in the seagrass meadow dominated by H. beccarii and the mangrove forest dominated by A. marina, with microplastics enriched by 1.3 to 17.6 times compared to their corresponding unvegetated sites. The abundance of microplastics varied greatly from 17.68 ± 8.10 to 611.75 ± 81.52 particles per kg of dry sediment, with the highest abundance in A. marina mangrove sediments. A strong positive correlation was found between the abundance of microplastics and the particulate organic carbon content at all study sites (Pearson, R = 0.86, p < 0.01). Higher diversity of microplastic colors and size was found in the H. beccarii meadow, and higher diversity of shapes was found in the A. marina forest. Our results added new insights to the understanding of the mechanism of microplastic trapping by BCEs and coupled the behavior of microplastics with the organic carbon in the sediment.

Bioturbation may not always enhance the metabolic capacity of organic polluted sediments.

Marine sediments are a major sink of organic matter, playing a crucial role in the global cycling of major elements. Macrofauna, through the reworking of particles and movement of solutes (bioturbation), enhances oxic conditions and the sediment metabolic capacity. Increases in the inputs of organic matter can lead to profound changes in the seabed and impact benthic ecological functions. Through a microcosm experiment, the effect of bioturbation of the polychaete Lumbrineris latreilli on biogeochemical fluxes under scenarios of increasing loads of organic matter was quantified. We found that bioturbation can buffer the negative consequences of anoxic conditions produced by organic enrichment, preventing the build-up of toxic by-products derived from anaerobic metabolic pathways by maintaining oxic conditions. However, the maintenance of oxic conditions by bioturbation is at the expense of limiting the sediment metabolic capacity. The maintenance of oxic conditions may limit anaerobic metabolic pathways, and consequently, the metabolic capacity of sediment. Thus, under organic matter pollution conditions, bioturbation may lessen the metabolic capacity of the sediment.

Root microbiomes as indicators of seagrass health.

The development of early warning indicators that identify ecosystem stress is a priority for improving ecosystem management. As microbial communities respond rapidly to environmental disturbance, monitoring their composition could prove one such early indicator of environmental stress. We combined 16S rRNA gene sequencing of the seagrass root microbiome of Halophila ovalis with seagrass health metrics (biomass, productivity and Fsulphide) to develop microbial indicators for seagrass condition across the Swan-Canning Estuary and the Leschenault Estuary (south-west Western Australia); the former had experienced an unseasonal rainfall event leading to declines in seagrass health. Microbial indicators detected sites of potential stress that other seagrass health metrics failed to detect. Genera that were more abundant in 'healthy' seagrasses included putative methylotrophic bacteria (e.g. Methylotenera and Methylophaga), iron cycling bacteria (e.g. Deferrisoma and Geothermobacter) and N2 fixing bacteria (e.g. Rhizobium). Conversely, genera that were more abundant in 'stressed' seagrasses were dominated by putative sulphur-cycling bacteria, both sulphide-oxidising (e.g. Candidatus Thiodiazotropha and Candidatus Electrothrix) and sulphate-reducing (e.g. SEEP-SRB1, Desulfomonile and Desulfonema). The sensitivity of the microbial indicators developed here highlights their potential to be further developed for use in adaptive seagrass management, and emphasises their capacity to be effective early warning indicators of stress.

Seagrass beds acting as a trap of microplastics - Emerging hotspot in the coastal region?

Microplastics is an emerging environmental problem in the world. However, presence and fate of microplastics in seagrass meadows are barely known. In this study, the abundance and diversity of microplastic from Enhalus acodoides vegetated sites and bare sites were quantified and characterized in Xincun bay and Li'an bay, Hainan, China. Microplastics ranged from 80.0 to 884.5 particles per kg of dry sediment, and fibers were the dominant shape. The most frequent colors of microplastics were blue, transparent and black. The dominant size of microplastics was in the range of 125-250 µm. And the seagrass sediments were enriched in microplastics 2.1 and 2.9 times for Xincun bay and Li'an bay, respectively. The trap effect of seagrass was non-selective regarding the shape, color and size of microplastics. High anthropogenic pollution and poor beach management may contribute to higher concentrations of microplastics in Li'an bay.

Variation of carbon contents in eelgrass ( Zostera marina) sediments implied from depth profiles.

Seagrass meadows are able to store significant amounts of organic carbon in their underlying sediment, but global estimates are uncertain partly owing to spatio-temporal heterogeneity between and within areas and species. In order to provide robust estimates, there is a need to better understand the fate of, and mechanisms behind, organic carbon storage. In this observational study, we analyse a suite of biotic and abiotic parameters in sediment cores from 47 different eelgrass ( Zostera marina) beds spanning the distributional range of the Northern Hemisphere. Depth profiles of particulate organic carbon (POC) revealed three patterns of vertical distribution where POC either increased, decreased or showed no pattern with sediment depth. These categories exhibited distinct profiles of δ13C and C:N ratios, where high POC profiles had a proportionally larger storage of eelgrass-derived material whereas low POC profiles were dominated by phytoplanktonic and macroalgal material. However, high POC did not always translate into high carbon density. Nevertheless, this large-scale dataset provides evidence that the variability in organic matter source in response to natural and anthropogenic environmental changes affects the potential role of eelgrass beds as POC sinks, particularly where eelgrass decline is observed.

Screening of seaweeds in the East China Sea as potential bio-monitors of heavy metals.

Seaweeds are good bio-monitors of heavy metal pollution and have been included in European coastal monitoring programs. However, data for seaweed species in China are scarce or missing. In this study, we explored the potential of seaweeds as bio-monitor by screening the natural occurring seaweeds in the "Kingdom of seaweed and shellfish" at Dongtou Islands, the East China Sea. Totally, 12 seaweed species were collected from six sites, with richness following the sequence of Rhodophyta > Phaeophyta > Chlorophyta. The concentration of heavy metals (Cu, Cr, Ni, Zn, Pb, Cd, As) in the seaweeds was determined, and the bioaccumulation coefficient was calculated. A combination of four seaweeds, Pachydictyon coriaceum, Gelidium divaricatum, Sargassum thunbergii, and Pterocladiella capillacea, were proposed as bio-monitors due to their high bioaccumulation capabilities of specific heavy metals in the East China Sea and hence hinted the importance of using seaweed community for monitoring of pollution rather than single species. Our results provide first-hand data for the selection of bio-monitor species for heavy metals in the East China Sea and contribute to selection of cosmopolitan bio-monitor communities over geographical large area, which will benefit the establishment of monitoring programs for coastal heavy metal contamination.

Species-specific response to sulfide intrusion in native and exotic Mediterranean seagrasses under stress.

We explored the sulfur dynamics and the relationships between sediment sulfur and nutrient pools, seagrass structural and physiological variables and sulfide intrusion in native (Posidonia oceanica, Cymodocea nodosa) and exotic (Halophila stipulacea) Mediterranean seagrasses at six sites affected by cumulative anthropogenic pressures to understand the factors controlling sulfide intrusion in seagrass. Sensitive indicators of seagrass stress (leaf TN, δ15N, TS, Fsulfide) were increased at several sites, implying that seagrasses are under pressure. Sulfide intrusion was not related to sediment TOC but it was negatively related to shoot size and below-ground biomass. Sulfide intrusion in seagrass tissue was high in P. oceanica (12-17%) and considerably higher in C. nodosa (27-35%). Intrusion was particularly high in H. stipulacea (30-50%), suggesting that its possible biogeographical expansion due to warming of the Mediterranean may result in accumulation of sulfides in the sediments and hypoxia/anoxia with further implications in ecosystem function.

Metabolomics Reveals Cryptic Interactive Effects of Species Interactions and Environmental Stress on Nitrogen and Sulfur Metabolism in Seagrass.

Eutrophication of estuaries and coastal seas is accelerating, increasing light stress on subtidal marine plants and changing their interactions with other species. To date, we have limited understanding of how such variations in environmental and biological stress modify the impact of interactions among foundational species and eventually affect ecosystem health. Here, we used metabolomics to assess the impact of light reductions on interactions between the seagrass Zostera marina, an important habitat-forming marine plant, and the abundant and commercially important blue mussel Mytilus edulis. Plant performance varied with light availability but was unaffected by the presence of mussels. Metabolomic analysis, on the other hand, revealed an interaction between light availability and presence of M. edulis on seagrass metabolism. Under high light, mussels stimulated seagrass nitrogen and energy metabolism. Conversely, in low light mussels impeded nitrogen and energy metabolism, and enhanced responses against sulfide toxicity, causing inhibited oxidative energy metabolism and tissue degradation. Metabolomic analysis thereby revealed cryptic changes to seagrass condition that could not be detected by traditional approaches. Our findings suggest that coastal eutrophication and associated reductions in light may shift seagrass-bivalve interactions from mutualistic to antagonistic, which is important for conservation management of seagrass meadows.

Eelgrass (Zostera marina) Food Web Structure in Different Environmental Settings.

This study compares the structure of eelgrass (Zostera marina L.) meadows and associated food webs in two eelgrass habitats in Denmark, differing in exposure, connection to the open sea, nutrient enrichment and water transparency. Meadow structure strongly reflected the environmental conditions in each habitat. The eutrophicated, protected site had higher biomass of filamentous algae, lower eelgrass biomass and shoot density, longer and narrower leaves, and higher above to below ground biomass ratio compared to the less nutrient-enriched and more exposed site. The faunal community composition and food web structure also differed markedly between sites with the eutrophicated, enclosed site having higher biomass of consumers and less complex food web. These relationships resulted in a column shaped biomass distribution of the consumers at the eutrophicated site whereas the less nutrient-rich site showed a pyramidal biomass distribution of consumers coupled with a more diverse consumer community. The differences in meadow and food web structure of the two seagrass habitats, suggest how physical setting may shape ecosystem response and resilience to anthropogenic pressure. We encourage larger, replicated studies to further disentangle the effects of different environmental variables on seagrass food web structure.

Sulfide Intrusion and Detoxification in the Seagrass Zostera marina.

Gaseous sulfide intrusion into seagrasses growing in sulfidic sediments causes little or no harm to the plant, indicating the presence of an unknown sulfide tolerance or detoxification mechanism. We assessed such mechanism in the seagrass Zostera marina in the laboratory and in the field with scanning electron microscopy coupled to energy dispersive X-ray spectroscopy, chromatographic and spectrophotometric methods, and stable isotope tracing coupled with a mass balance of sulfur compounds. We found that Z. marina detoxified gaseous sediment-derived sulfide through incorporation and that most of the detoxification occurred in underground tissues, where sulfide intrusion was greatest. Elemental sulfur was a major detoxification compound, precipitating on the inner wall of the aerenchyma of underground tissues. Sulfide was metabolized into thiols and entered the plant sulfur metabolism as well as being stored as sulfate throughout the plant. We conclude that avoidance of sulfide exposure by reoxidation of sulfide in the rhizosphere or aerenchyma and tolerance of sulfide intrusion by incorporation of sulfur in the plant are likely major survival strategies of seagrasses in sulfidic sediments.

Impact of pond aquaculture effluents on seagrass performance in NE Hainan, tropical China.

The impact of pond aquaculture effluents on the distribution and performance of seagrasses was examined in NE Hainan, tropical China. Samples were taken along transects in three back-reef areas with different extent of aquaculture production in their hinterland. High δ(15)N in seagrass leaves and epiphytes (6-9‰) similar to values in pond effluents documented aquaculture as dominant nitrogen source in the back-reefs with decreasing impact with distance from shore. Seagrass species abundance, shoot density and biomass were lower and concentrations of nutrients, chlorophyll and suspended matter were higher at nearshore sites with high and moderate pond abundance than at the control site. High epiphyte loads and low δ(34)S in seagrass leaves suggest temporal shading and sulphide poisoning of the nearshore seagrasses. Observed gradients in environmental parameters and seagrass performance indicate that the distance from the pond outlets and size of the adjacent pond agglomeration are major determinants of seagrass degradation.

Mussels as a tool for mitigation of nutrients in the marine environment.

Long-line mussel farming has been proposed as a mitigation tool for removal of excess nutrients in eutrophic coastal waters. A full-scale mussel farm optimized for cost efficient nutrient removal was established in the eutrophic Skive Fjord, Denmark where biological and economic parameters related to nutrient removal was monitored throughout a full production cycle (1 yr). The results showed that it was possible to obtain a high area specific biomass of 60 t WW ha(-1) eqvivalent to a nitrogen and phosphorus removal of 0.6-0.9 and 0.03-0.04 t ha(-1)yr, respectively. The analysis of the costs related to establishment, maintenance and harvest revealed that mussel production optimized for mitigation can be carried out at a lower cost compared to mussel production for (human) consumption. The costs for nutrient removal was 14.8 € kg(-1)N making mitigation mussel production a cost-efficient measure compared to the most expensive land-based measures.

Global warming enhances sulphide stress in a key seagrass species (NW Mediterranean).

The build-up of sulphide concentrations in sediments, resulting from high inputs of organic matter and the mineralization through sulphate reduction, can be lethal to the benthos. Sulphate reduction is temperature dependent, thus global warming may contribute to even higher sulphide concentrations and benthos mortality. The seagrass Posidonia oceanica is very sensitive to sulphide stress. Hence, if concentrations build up with global warming, this key Mediterranean species could be seriously endangered. An 8-year monitoring of daily seawater temperature, the sulphur isotopic signatures of water (δ(34)S(water)), sediment (δ(34)SCRS ) and P. oceanica leaf tissue (δ(34)S(leaves)), along with total sulphur in leaves (TS(leaves)) and annual net population growth along the coast of the Balearic archipelago (Western Mediterranean) allowed us to determine if warming triggers P. oceanica sulphide stress and constrains seagrass survival. From the isotopic S signatures, we estimated sulphide intrusion into the leaves (F(sulphide)) and sulphur incorporation into the leaves from sedimentary sulphides (SS(leaves)). We observed lower δ(34)S(leaves), higher F(sulphide) and SS(leaves) coinciding with a 6-year period when two heat waves were recorded. Warming triggered sulphide stress as evidenced by the negative temperature dependence of δ(34)S(leaves) and the positive one of F(sulphide), TS(leaves) and SS(leaves). Lower P. oceanica net population growth rates were directly related to higher contents of TS(leaves). At equivalent annual maximum sea surface water temperature (SST(max)), deep meadows were less affected by sulphide intrusion than shallow ones. Thus, water depth acts as a protecting mechanism against sulphide intrusion. However, water depth would be insufficient to buffer seagrass sulphide stress triggered by Mediterranean seawater summer temperatures projected for the end of the 21st century even under scenarios of moderate greenhouse gas emissions, A1B. Mediterranean warming, therefore, is expected to enhance P. oceanica sulphide stress, and thus compromise the survival of this key habitat along its entire depth distribution range.

Biogeochemical malfunctioning in sediments beneath a deep-water fish farm.

We investigated the environmental impact of a deep water fish farm (190 m). Despite deep water and low water currents, sediments underneath the farm were heavily enriched with organic matter, resulting in stimulated biogeochemical cycling. During the first 7 months of the production cycle benthic fluxes were stimulated >29 times for CO(2) and O(2) and >2000 times for NH(4)(+), when compared to the reference site. During the final 11 months, however, benthic fluxes decreased despite increasing sedimentation. Investigations of microbial mineralization revealed that the sediment metabolic capacity was exceeded, which resulted in inhibited microbial mineralization due to negative feed-backs from accumulation of various solutes in pore water. Conclusions are that (1) deep water sediments at 8 °C can metabolize fish farm waste corresponding to 407 and 29 mmol m(-2) d(-1) POC and TN, respectively, and (2) siting fish farms at deep water sites is not a universal solution for reducing benthic impacts.

A meta-analysis of seaweed impacts on seagrasses: generalities and knowledge gaps.

Seagrasses are important habitat-formers and ecosystem engineers that are under threat from bloom-forming seaweeds. These seaweeds have been suggested to outcompete the seagrasses, particularly when facilitated by eutrophication, causing regime shifts where green meadows and clear waters are replaced with unstable sediments, turbid waters, hypoxia, and poor habitat conditions for fishes and invertebrates. Understanding the situations under which seaweeds impact seagrasses on local patch scales can help proactive management and prevent losses at greater scales. Here, we provide a quantitative review of available published manipulative experiments (all conducted at the patch-scale), to test which attributes of seaweeds and seagrasses (e.g., their abundances, sizes, morphology, taxonomy, attachment type, or origin) influence impacts. Weighted and unweighted meta-analyses (Hedges d metric) of 59 experiments showed generally high variability in attribute-impact relationships. Our main significant findings were that (a) abundant seaweeds had stronger negative impacts on seagrasses than sparse seaweeds, (b) unattached and epiphytic seaweeds had stronger impacts than 'rooted' seaweeds, and (c) small seagrass species were more susceptible than larger species. Findings (a) and (c) were rather intuitive. It was more surprising that 'rooted' seaweeds had comparatively small impacts, particularly given that this category included the infamous invasive Caulerpa species. This result may reflect that seaweed biomass and/or shading and metabolic by-products like anoxia and sulphides could be lower for rooted seaweeds. In conclusion, our results represent simple and robust first-order generalities about seaweed impacts on seagrasses. This review also documented a limited number of primary studies. We therefore identified major knowledge gaps that need to be addressed before general predictive models on seaweed-seagrass interactions can be build, in order to effectively protect seagrass habitats from detrimental competition from seaweeds.

Effect of temperature on biogeochemistry of marine organic-enriched systems: implications in a global warming scenario.

Coastal biogeochemical cycles are expected to be affected by global warming. By means of a mesocosm experiment, the effect of increased water temperature on the biogeochemical cycles of coastal sediments affected by organic-matter enrichment was tested, focusing on the carbon, sulfur, and iron cycles. Nereis diversicolor was used as a model species to simulate macrofaunal bioirrigation activity in natural sediments. Although bioirrigation rates of N. diversicolor were not temperature dependent, temperature did have a major effect on the sediment metabolism. Under organic-enrichment conditions, the increase in sediment metabolism was greater than expected and occurred through the enhancement of anaerobic metabolic pathway rates, mainly sulfate reduction. There was a twofold increase in sediment metabolism and the accumulation of reduced sulfur. The increase in the benthic metabolism was maintained by the supply of electron acceptors through bioirrigation and as a result of the availability of iron in the sediment. As long as the sediment buffering capacity toward sulfides is not surpassed, an increase in temperature might promote the recovery of organic-enriched sediments by decreasing the time for mineralization of excess organic matter.