Sulfide Intrusion Research Articles

Reduced seagrass resilience due to environmental and anthropogenic effects may lead to future die-off events in Florida Bay

Coastal systems often depend on foundation species such as seagrasses that are supported by self-facilitation. Seagrass meadows, however, are threatened worldwide due to climate change and local human impact, disrupting self-facilitation leading to system instability. Florida Bay is a large seagrass dominated coastal ecosystem that suffered from multiple seagrass mortality events over the last half century driven by hypoxia, high water temperatures, hypersalinity, and high biological oxygen demand. These conditions reduce the amount of photosynthetically-derived oxygen in the plant causing sulphide intrusion into meristematic tissues. Using a bay-wide sampling design and long-term monitoring trends of seagrass condition, we investigated the current state of the meadows, sediment characteristics (e.g., organic matter, sulphide, nutrients) and discuss how climate stressors interact with plant and sediment oxygen dynamics. Our survey revealed that at sites where seagrass had been previously denuded by die-off, the dominant seagrass Thalassia testudinum had not recovered, while the pioneering seagrass Halodule wrightii recolonized the impacted areas. Organic matter and sulphide levels were higher at the impacted sites, apparently a persistent characteristic of the formerly dense T. testudinum meadows in central and western Florida Bay. These sediment conditions promote sulphide intrusion of T. testudinum belowground tissue under anoxic conditions. Plant oxidation initially buffers sulphide intrusion, but disruption of this oxidation mechanism due to changing environmental conditions results in widespread mortality and seagrass community collapse. Climate change cannot be fully mitigated by local management, however, attempts can be made to control critical salinity and oxygen levels by increasing freshwater input, reducing hypersalinity and aiming to keep the internal seagrass oxidation feedback intact. Our study shows that the Florida Bay seagrass ecosystem is still recovering four years post die-off and continues to be susceptible to future climate change and system degradation.

Spatial and Temporal Variation of C, N, and S Stable Isotopes and Seagrass Coverage Related to Eutrophication Stress in Zostera marina

Zostera marina is an ecologically valuable species that has been declining due to anthropogenic environmental stressors. In this study, spatial and temporal indicators of eelgrass stress, such as coverage and biomass, were compared with the isotopic composition of C, N, and S to understand the mechanism(s) of plant stress. Eelgrass samples were collected in June, July, and August of 2020 at five stations along an estuary spatial gradient in the southern Gulf of St. Lawrence to measure above- and below-ground biomass and tissue isotopes in eelgrass leaves and roots/rhizomes. Eelgrass biomass was lowest at the innermost sampling station, which coincided with eutrophication-induced hypoxia relative to outer sampling stations. δ13C levels at the upstream station were depleted compared to downstream stations. Comparatively, δ15N and δ34S findings were not correlated with plant biomass. Thus, sulfide intrusion was not a major stressor for eelgrass in this estuary. Between the years 2014 and 2020, eelgrass coverage was found to have increased, which coincided with high and low recorded external nutrient loads from the Wheatley River, respectively. Ultimately, these findings indicate that isotopic composition and biomass can be useful in assessing the health of eelgrass in temperate estuaries.

Combined exposure to CO2 and H2S significantly reduces the performance of the Mediterranean seagrass Posidonia oceanica: Evidence from a volcanic vent

AbstractAlthough seagrasses are expected to thrive in future acidified oceans by overcoming low CO2 diffusion into plant tissues, the co‐occurrence of environmental stressors may affect their growth. Volcanic CO2 vents are often associated with toxic gases and metal‐rich fluids representing ideal sites to assess the effects of multiple stressors. We evaluated the response of Posidonia oceanica growing near shallow CO2 vents characterized by H2S spill‐out by comparing meadow structure and phenology to an area with no gas emissions. Seagrass descriptors at meadow, shoot and leaf level indicated that P. oceanica experienced stressful conditions at the vent area, in clear contrast to the flourishing features of P. oceanica previously described at CO2 vents with no evidence of toxic inputs. Furthermore, the reduction in both leaf δ34S and growth at the vent area indicates that sulfide intrusion occurs and affects seagrass growth performance, dampening the expected beneficial effects of high CO2 levels.

The barrier to radial oxygen loss protects roots against hydrogen sulphide intrusion and its toxic effect.

The root barrier to radial O2 loss (ROL) is a key root trait preventing O2 loss from roots to anoxic soils thereby enabling root growth into anoxic, flooded soils. We hypothesized that the ROL barrier can also prevent intrusion of hydrogen sulphide (H2 S), a potent phytotoxin in flooded soils. Using H2 S and O2 sensitive microsensors, we measured the apparent permeance to H2 S of rice roots, tested whether restricted H2 S intrusion reduced its adverse effects on root respiration, and if H2 S could induce the formation of a ROL barrier. The ROL barrier reduced apparent permeance to H2 S by almost 99%, greatly restricting H2 S intrusion. The ROL barrier acted as a shield towards H2 S; O2 consumption in roots with a ROL barrier remained unaffected at high H2 S concentration (500 μM), compared to a 67% decline in roots without a barrier. Importantly, low H2 S concentrations induced the formation of a ROL barrier. In conclusion, the ROL barrier plays a key role in protecting against H2 S intrusion, and H2 S can act as an environmental signalling molecule for the induction of the barrier. The study demonstrates the multiple functions of the suberized/lignified outer part of the rice root beyond that of restricting ROL.

Sulfide intrusion in a habitat forming seagrass can be predicted from relative abundance of sulfur cycling genes in sediments

Sulfide intrusion from sediments is an increasingly recognized contributor to seagrass declines globally, yet the relationship between sediment microorganisms and sulfide intrusion has received little attention. Here, we use metagenomic sequencing and stable isotope (34S) analysis to examine this relationship in Cockburn Sound, Australia, a seagrass-dominated embayment with a gradient of sulfide stress and seagrass declines. There was a significant positive relationship between sulfide intrusion into seagrasses and sulfate reduction genes in sediment microbial communities, which was greatest at sites with long term seagrass declines. This is the first demonstration of a significant link between sulfur cycling genes present in seagrass sediments and sulfide intrusion in a habitat-forming seagrass that is experiencing long-term shoot density decline. Given that microorganisms respond rapidly to environmental change, the quantitative links established in this study can be used as a potential management tool to enable the prediction of sulfide stress on large habitat forming seagrasses; a global issue expected to worsen with climate change.

Sand supplementation favors tropical seagrass Thalassia hemprichii in eutrophic bay: implications for seagrass restoration and management

BackgroundSediment is crucial for the unique marine angiosperm seagrass growth and successful restoration. Sediment modification induced by eutrophication also exacerbates seagrass decline and reduces plantation and transplantation survival rates. However, we lack information regarding the influence of sediment on seagrass photosynthesis and the metabolics, especially regarding the key secondary metabolic flavone. Meanwhile, sulfation of flavonoids in seagrass may mitigate sulfide intrusion, but limited evidence is available.ResultsWe cultured the seagrass Thalassia hemprichii under controlled laboratory conditions in three sediment types by combining different ratios of in-situ eutrophic sediment and coarse beach sand. We examined the effects of beach sand mixed with natural eutrophic sediments on seagrass using photobiology, metabolomics and isotope labelling approaches. Seagrasses grown in eutrophic sediments mixed with beach sand exhibited significantly higher photosynthetic activity, with a larger relative maximum electron transport rate and minimum saturating irradiance. Simultaneously, considerably greater belowground amino acid and flavonoid concentrations were observed to counteract anoxic stress in eutrophic sediments without mixed beach sand. This led to more positive belowground stable sulfur isotope ratios in eutrophic sediments with a lower Eh.ConclusionsThese results indicated that coarse beach sand indirectly enhanced photosynthesis in T. hemprichii by reducing sulfide intrusion with lower amino acid and flavonoid concentrations. This could explain why T. hemprichii often grows better on coarse sand substrates. Therefore, it is imperative to consider adding beach sand to sediments to improve the environmental conditions for seagrass and restore seagrass in eutrophic ecosystems.

Low irradiance disrupts the internal O2 dynamics of seagrass (Thalassia testudinum) leading to shoot meristem H2S intrusion

Hypoxia and hydrogen sulfide (H2S) intrusion at night contribute to large-scale seagrass mortality events world-wide. Declining water quality has lowered irradiance and enhanced hypoxia in seagrass ecosystems, but linkages between low irradiance and seagrass internal pO2in situ are not well understood. We examined low irradiance effects on leaf and meristem pO2 dynamics of a dominant tropical seagrass, Thalassia testudinum, using microsensors over multiple diurnal cycles. Further, we determined how O2 dynamics affect H2S intrusion into shoot meristems under low irradiance. Sequential days of low irradiance disrupted internal leaf O2 status in three ways: i) causing a longer lag in morning O2 pressurization, ii) depressing maximum pO2 during the day, and iii) shortening the time where maximum pO2 was sustained into the afternoon. There was a close relationship between leaf and water column pO2 during the day (R2 = 0.93 ± 0.09); thus, internal O2 dynamics appear more dependent on water column pO2 under low irradiance. The nighttime minimum of leaf and meristem pO2 (1.2 and 0.4 kPa, respectively) were very low and the length of time the meristem sustained hypoxia (< 1.5 kPa pO2) at night was high (9:40 h:min). H2S intrusion into the meristem at night following 24–48 h of shading was persistent, likely the combined effect of limited internal O2 flow without saturating irradiances for photosynthesis, and a breakdown of the sediment oxic microshield. Under low irradiance, more frequent intrusions of H2S to the meristems increases the chances of large-scale seagrass mortality events at night.

Variation in sediment and seagrass characteristics reflect multiple stressors along a nitrogen‐enrichment gradient in a New England lagoon

AbstractWe examined concentrations of organic carbon, dissolved sulfides, total sediment sulfur, and stable sulfur isotope ratios in seagrass leaf tissues across a nitrogen‐enrichment gradient in a coastal marine ecosystem (Cape Cod, Massachusetts) in 2007–2010 and 2017–2019. We also measured seagrass aboveground and belowground biomass, epibiota biomass, and leaf chlorophyll content. Seagrasses were present at all sites in the former period but were lost at our most nitrogen‐impacted site (Snug Harbor) by 2011. In 2007–2010, sediment organic carbon and dissolved sulfides were highest in Snug Harbor and decreased along the gradient; leaf tissues depleted in 34S also indicated higher sulfide intrusion into seagrass tissues in more eutrophic areas. By 2017–2019, sediment organic carbon and pore‐water soluble sulfides had decreased in Snug Harbor, but had increased at the intermediate site, to levels found at the most impacted site prior to the seagrass die‐off. Again, leaf tissue 34S depletion reflected this pattern, indicating seagrasses were exposed to the highest sulfides at the intermediate site. The decreases in sediment organic carbon and soluble sulfides in Snug Harbor years after the loss of the seagrasses illustrate a feedback between high organic matter in seagrass beds and increasing stressors like elevated soluble sulfides in nutrient‐enriched systems. We found significant relationships between sediment conditions and seagrass responses, including greater aboveground to belowground biomass ratios, epibiota biomass, and 34S‐depleted leaves at sites with high pore‐water sulfide and highly organic sediments. Our research suggests that the reduction of anthropogenic nitrogen entering the harbor is necessary for improving sediment quality and preventing seagrass mortality.

An assessment of hydrogen sulfide intrusion in the seagrass Halodule wrightii

Abstract Hydrogen sulfide (H2S, “sulfide”) is a naturally occurring component of the marine sediment. Eutrophication of coastal waters, however, can lead to an excess of sulfide production that can prove toxic to seagrasses. We used stable sulfur isotope ratio (δ34S) measurements to assess sulfide intrusion in the seagrass Halodule wrightii, a semi-tropical species found throughout the Gulf of Mexico, Caribbean Sea, and both western and eastern Atlantic coasts. We found a gradient in δ34S values (−5.58 ± 0.54‰+13.58 ± 0.30‰) from roots to leaves, in accordance with prior observations and those from other species. The results may also represent the first values reported for H. wrightii rhizome tissue. The presence of sulfide-derived sulfur in varying proportions (15–55%) among leaf, rhizome, and root tissues suggests H. wrightii is able to assimilate sedimentary H2S into non-toxic forms that constitute a significant portion of the plant’s total sulfur content.

Landscape-scale variation in a sulfur-based sediment stress indicator for the seagrass Thalassia testudinum in Florida Bay, USA

Intrusion of sediment-derived hydrogen sulfide into above-ground tissues of seagrasses is correlated with reduced growth and has been linked to large-scale die-offs of Thalassia testudinum in Florida Bay, USA. In May of 2019, leaves from T. testudinum short shoots at 350 sites within 13 basins across Florida Bay were collected to investigate bay-wide variation in a novel, stable sulfur isotope-based indicator of hydrogen sulfide intrusion [(δ34Sleaf + 30)/total sulfur content]. The components of this sediment stress indicator (SSI), total sulfur content (% dry weight) and δ34S, were negatively correlated (R2 = -0.24), indicating greater sediment sulfide exposure in plants with higher sulfur content. Generalized additive model selection revealed that SSI was best predicted by a model which included T. testudinum cover, sediment depth, and Halodule wrightii cover (R2 = 0.24, weight = 0.48). Macrophyte communities dominated by dense T. testudinum climax communities and with deep sediments, which are characteristics associated with die-off, had the lowest SSI values, indicating greatest sulfide intrusion. Sites within the area of a recent (2015) seagrass die-off had significantly higher SSI values than nearby, non-die-off-affected sites (mean ± SE: 44.1 ± 1.5 vs. 40.2 ± 1.6, Wilcoxon p &lt; 0.05). The recent die-off sites also had lower T. testudinum cover and higher H. wrightii cover than the non-die-off sites, indicating they were in the midst of recovery and at an early successional stage. Our findings indicate that SSI may provide a minimally destructive indicator of chronic sulfide intrusion in T. testudinum in Florida Bay.

First Field-Based Evidence That the Seagrass-Lucinid Mutualism Can Mitigate Sulfide Stress in Seagrasses

Seagrass meadows form vital ecological components of coastal zones worldwide, but are rapidly declining. Large-scale seagrass diebacks have been related to accumulation of toxic sulfide in the sediment, a phenomenon predicted to occur more frequently in the near future due to ongoing global warming and increasing organic loading of coastal systems worldwide. Recently, a facultative mutualism between seagrasses and lucinid bivalves with endosymbiotic sulfide-consuming gill bacteria was discovered that may prevent toxic sulfide accumulation in seagrass sediments. Yet, direct field-based evidence for the importance of this mutualism in alleviating sulfide stress in seagrasses is currently lacking, as well as how its role may change when sediment sulfide levels increase due to environmental change. Here, we investigated the sulfide detoxification function of this seagrass-lucinid mutualism and its resilience to organic-loading induced sulfide stress in a temperate lagoon system (Thau lagoon, France), using a correlative field survey and a full factorial field experiment. The field survey revealed a strong positive correlation between seagrass above-ground biomass and lucinid densities, and pore water sulfide concentrations close to zero at all sites. Furthermore, the field experiment revealed that addition of organic matter (starch mixed with sucrose) increased sedimentary sulfide intrusion into seagrass (Zostera noltei) leaves (a proxy for sulfide stress), while experimentally enhanced lucinid densities reduced sulfide intrusion, regardless of addition of organic matter. Moreover, addition of organic matter reduced seagrass rhizome biomass and increased pore water sulfide levels, lucinid tissue sulfur content, lucinid condition (expressed as flesh/shell dry weight ratio), and total lucinid biomass, while enhancement of lucinid densities reduced lucinid condition. These results provide the first field-based evidence that lucinid bivalves and their sulfide-oxidizing gill symbionts mitigate sulfide stress in seagrasses, and suggests that the dependence of seagrass on this seagrass-lucinid mutualism will increase under conditions of enhanced sediment sulfide production, as predicted for the near future. Therefore, we suggest that awareness of the ecological importance of the seagrass-lucinid mutualism may be instrumental for designing new measures for improving long-term restoration success and seagrass resilience to global change.

Oxygen loss from seagrass roots coincides with colonisation of sulphide-oxidising cable bacteria and reduces sulphide stress.

Seagrasses thrive in anoxic sediments where sulphide can accumulate to phytotoxic levels. So how do seagrasses persist in this environment? Here, we propose that radial oxygen loss (ROL) from actively growing root tips protects seagrasses from sulphide intrusion not only by abiotically oxidising sulphides in the rhizosphere of young roots, but also by influencing the abundance and spatial distribution of sulphate-reducing and sulphide-oxidising bacteria. We used a novel multifaceted approach combining imaging techniques (confocal fluorescence in situ hybridisation, oxygen planar optodes, and sulphide diffusive gradients in thin films) with microbial community profiling to build a complete picture of the microenvironment of growing roots of the seagrasses Halophila ovalis and Zostera muelleri. ROL was restricted to young root tips, indicating that seagrasses will have limited ability to influence sulphide oxidation in bulk sediments. On the microscale, however, ROL corresponded with decreased abundance of potential sulphate-reducing bacteria and decreased sulphide concentrations in the rhizosphere surrounding young roots. Furthermore, roots leaking oxygen had a higher abundance of sulphide-oxidising cable bacteria; which is the first direct observation of these bacteria on seagrass roots. Thus, ROL may enhance both abiotic and bacterial sulphide oxidation and restrict bacterial sulphide production around vulnerable roots, thereby helping seagrasses to colonise sulphide-rich anoxic sediments.

Local Regime Shifts Prevent Natural Recovery and Restoration of Lost Eelgrass Beds Along the Swedish West Coast

Along the Swedish northwest coast, over 60% of the eelgrass meadows have been lost since the 1980s. Despite improved water quality, no recovery has occurred, and restoration is presently considered to mitigate historical losses. However, the factors preventing natural recovery of eelgrass are not known, and it is not clear if conditions would allow restoration. Here, we present the results from 5 years of field studies with the aim of identifying the key processes affecting eelgrass growth and survival at historical eelgrass areas. Continuous light measurements and comparison with historic eelgrass distribution indicate that maximum depth distribution has decreased locally with 1.5–2.3 m in areas that have lost large eelgrass beds in the last 10–30 years. Field studies suggest that wind-driven local resuspension of sediments that are no longer stabilized by eelgrass beds is the main cause behind the deteriorated light conditions. Field experiments show that a combination of low light condition and disturbance from drifting algal mats prevents eelgrass recovery in these areas, whereas the sulfide intrusion from the sediment and dislodgement of shoots by waves had little effect on growth and survival. These results suggest that local regime shifts acting on a scale of 40–200 ha have occurred after the loss of eelgrass beds, where increased sediment resuspension and proliferation of drifting algal mats act as feedback mechanisms that prevent both natural recovery and restoration of eelgrass. The feedbacks appear to be interacting and causing an accelerating loss of eelgrass that is presently spreading to neighboring areas.

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.

Belowground stressors and long-term seagrass declines in a historically degraded seagrass ecosystem after improved water quality

Continued seagrass declines in ecosystems with improved water quality may be driven by sediment stressors. One of the most cited examples of a seagrass ecosystem with declines is Cockburn Sound, Western Australia, where 75% of seagrasses (2169 ha) were lost in the 1960s–1980s due to poor water quality. Water quality has subsequently improved in Cockburn Sound, yet shoot density declines continue in some areas. Here, we investigated if sediment stressors (sulfide intrusion and heavy metals) contributed to declining Posidonia sinuosa shoot densities in Cockburn Sound. Seagrass δ34S were depleted at sites with a history of seagrass declines, indicating seagrasses at these sites were under sulfide stress. Heavy metals (Fe, Zn, Mn, Cr, Cu and Cd) in sediments and seagrasses did not show clear patterns with shoot density or biomass, and largely decreased from similar measurements in the late 1970s. However, seagrass cadmium concentrations were negatively correlated to seagrass biomass and shoot density. High cadmium concentrations interfere with sulfur metabolism in terrestrial plants, but impacts on seagrasses remain to be explored. Given that sulfide intrusion can prevent recolonization and drive seagrass declines, management plans in degraded seagrass ecosystems should include management of sediment stressors and water quality to provide comprehensive management.

Drivers of sulfide intrusion in Zostera muelleri in a moderately affected estuary in south-eastern Australia

The seagrass Zostera muelleri Irmisch ex Asch. is abundant in estuaries in Australia and is under pressure from coastal developments. We studied sulfide intrusion in Z. muelleri along a gradient of anthropogenic impact at five stations in the Wallis Lake estuary, Australia. Results showed differences in sediment biogeochemical conditions, seagrass metrics as well as nutrient content and sulfide intrusion along the gradient from the lower estuary (affected) to the lagoon (unaffected). Sulfide intrusion was driven by complex interactions and related to changes in seagrass morphology and sediment biogeochemistry and was modified by the exposure to wind and wave action. The sediments in the lower estuary had high contributions from phytoplanktonic detritus, whereas the organic pools in the lagoon were dominated by seagrass detritus. Despite high concentrations of organic matter, sulfide intrusion was lower at stations dominated by seagrass detritus, probably because of lower sulfide pressure from the less labile nature of organic matter. Porewater diffusive gradients in thin-film (DGT) sulfide samplers showed efficient sulfide reoxidation in the rhizosphere, with high sulfur incorporation in the plants from sedimentary sulfides being likely due to sulfate uptake from reoxidised sulfide. This is a unique adaptation of Z. muelleri, which allows high productivity in estuarine sediments.

Effects of point source of untreated sewage waters on seagrass (Zostera marina and Z. noltii) beds in the South-Western Black Sea

Tourist development is rapid on the South-West coast of the Black Sea, increasing the pressure on the coastal ecosystem. Zostera marina and Zostera noltii are abundant seagrasses in the Black Sea, but little is known about their sensitivity towards coastal developments. In this study plant nutrient content (N and P), sulphide intrusion, sediment organic matter, pore water pools of nutrients and sulphides, sulphate reduction rates as well as water column nutrients and chlorophyll-a concentrations were studied in a mixed seagrass meadow (Z. marina L. and Z. noltii Hornem.) located in an area receiving untreated sewage waters from a point source in Sozopol Bay, South-Western Black Sea. In addition, studies included population level metrics like density, biomass and shoot morphology. The point source impacted water quality in the area leading to local nutrient enrichment, development of filamentous macroalgae and reduced water transparency. Although sulphide intrusion was detected in Z. marina and thus indicating existence of sulphide pressure on the plants, there were no significant effects on the seagrass distribution except for Z. noltii being absent from the meadows close to the point source. The nutrient levels were low in the plants and in the sediments compared to global Z. marina and Z. noltii meadows, but the observed blooms of filamentous macroalgae, increase in sulphide intrusion and higher N contents of the plants during the peak tourist season are signs of eutrophication in Sozopol Bay most likely as a result of increased recreational activity on the coast.

Light indirectly mediates bivalve habitat modification and impacts on seagrass

Environmental context may influence the sign, strength, and mechanisms of species interactions but few studies have experimentally tested the potential for abiotic conditions to mediate interactions through multiple co-occurring stress pathways. Abiotic conditions may mediate species interactions by directly or indirectly influencing the effects of habitat-modifying organisms that are capable of simultaneously ameliorating and exacerbating multiple stressors. It was hypothesized that light availability changes seagrass metabolism and thereby indirectly regulates bivalve habitat modification and subsequent impacts on seagrasses by shifting net effects between alleviation of nutrient stress and intensification of sulfide stress. To test this hypothesis, manipulations of light availability and blue mussel (Mytilus edulis) abundance were made in eelgrass (Zostera marina) mesocosms and biogeochemical and plant responses were measured. Light modified the effect of mussels on porewater ammonium, but eelgrass was not nutrient limited and, therefore, mussels did not enhance growth. Mussels increased sediment sulfides irrespective of light availability and, by reducing net oxygen flux (production and respiration), mussels and low light availability exacerbated sulfide intrusion of eelgrass tissues. Surprisingly, sulfide stress did not affect plant growth, survival, or energy stores. Thus, habitat modification by mussels may represent a risk to eelgrass, especially during low productivity conditions, but eelgrass can resist harm from short-term stress, even during light limitation. These findings suggest that while small-scale bivalve impacts on seagrasses may be variable in oligotrophic estuaries, they have the potential to be negative in eutrophic systems, which are increasing globally.

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.

Sensitivity of Ruppia maritima and Zostera marina to sulfide exposure around roots

Ruppia maritima and Zostera marina often inhabit the same coastal areas and are often exposed to fairly high sulfide concentrations, with the former being most resilient. Our aim was to investigate differences in sulfide tolerance between the two species. Sensitivity towards sulfide in R. maritima and Z. marina was investigated in split chambers that allowed exposure of roots to a range of sulfide concentrations, while the leaves were kept under optimal light and oxygen conditions. Roots of R. maritima show a higher threshold towards sulfide (1.8–5.9mM H2S) than those of Z. marina (0–0.5mM H2S) after two weeks of incubation. The difference in sulfide sensitivity is attributed to different capacity of root oxygen leakage. Light and oxygen exposure enables oxygen transport from leaves down to the roots during both day and night. The oxic micro-shield around roots has a capacity to withstand sulfide intrusion until the sensitivity level is reached, when no other stressors are present. Leaves of exposed R. maritima plants are affected at similar sulfide concentrations as roots, while leaves of Z. marina are affected at higher sulfide concentrations (1–3.5mM H2S), reflecting different pathways of sulfide intrusion, root permeability and the ability to allocate energy away from exposed plant parts. Sensitivity towards sulfide occurs gradually and probably as a linear response to increasing sulfide concentration in R. maritima, but the exact pattern in Z. marina is still unclear. Other external stressors, such as anoxia, shading, or increased temperature may change the thresholds of the species by affecting radial oxygen loss from roots, and make especially R. maritima more prone to sulfide toxicity due to its highly permeable roots.