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Large nutrient levels and herbivory stress, particularly when acting together, drive a variety of responses in seagrass communities that ultimately may weaken their carbon balance. An in situ three-months experiment was carried out in two contrasting seasons to address the effects of two levels of nutrient load and three levels of artificial clipping on Cymodocea nodosa plants. Nutrient enrichment shifted the community from autotrophic to heterotrophic and reduced DOC fluxes in winter, whereas enhanced community carbon metabolism and DOC fluxes in summer. Herbivory stress decreased the net primary production in both seasons, whereas net DOC release increased in winter but decreased in summer. A reduction of seagrass food-web structure was observed under both disturbances evidencing impacts on the seagrass ecosystems services by altering the carbon transfer process and the loss of superficial OC, which may finally weaken the blue carbon storage capacity of these communities., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

Seagrass communities have been degraded worldwide experiencing elevated shoot density reduction by anthropogenic chronic pressures. This study aims to assess how a chronic (i.e., low intensity but long-lasting) impact that promotes reduced shoot density in a temperate seagrass population may affect community components and functioning. To this end, shoot density was reduced (0, 40, and 75 %) for three months in contrasting seasons (winter and summer), and assessed its effects on biotic components (i.e., seagrasses, macroalgae, macrofauna, and microphytobenthos), as well as on community carbon metabolism, dissolved organic carbon (DOC) fluxes and sediment organic matter (OM) content. Lower shoot densities enhanced the presence of macroalgae and microphytobenthos in the community, while macrofauna remained unchanged. Net community production was significantly reduced with the simulated reduction in shoot density in both seasons (up to 10-fold lower), which shifted the community in winter from being largely autotrophic (CO 2 sink) to heterotrophic (CO 2 source). This was due to the expected reduction in gross primary production, but also to the unexpected increase in community respiration (up to 2.2-fold higher). Since OM in the sediment was reduced in the simulated shoot density reduction treatments, the increase in sediment bacterial activity may help explain the increase in community respiration. DOC fluxes were also greatly reduced in both seasons (up to 5.5-fold lower), which coupled with the reduced net community production and loss of OM in the sediment may have a continued silent effect on blue carbon capture and storage capacity in this chronically stressed community. This study therefore highlights the importance of chronic impacts that promote the degradation of seagrass communities that may reduce their ability to provide highly valuable ecological services, including the ability to cope with the effects of climate change., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 The Authors. Published by Elsevier B.V. All rights reserved.)

Mangroves and saltmarshes are biogeochemical hotspots storing carbon in sediments and in the ocean following lateral carbon export (outwelling). Coastal seawater pH is modified by both uptake of anthropogenic carbon dioxide and natural biogeochemical processes, e.g., wetland inputs. Here, we investigate how mangroves and saltmarshes influence coastal carbonate chemistry and quantify the contribution of alkalinity and dissolved inorganic carbon (DIC) outwelling to blue carbon budgets. Observations from 45 mangroves and 16 saltmarshes worldwide revealed that >70% of intertidal wetlands export more DIC than alkalinity, potentially decreasing the pH of coastal waters. Porewater-derived DIC outwelling (81 ± 47 mmol m -2 d -1 in mangroves and 57 ± 104 mmol m -2 d -1 in saltmarshes) was the major term in blue carbon budgets. However, substantial amounts of fixed carbon remain unaccounted for. Concurrently, alkalinity outwelling was similar or higher than sediment carbon burial and is therefore a significant but often overlooked carbon sequestration mechanism., (© 2023. The Author(s).)

A crucial factor in the long-term survival of benthic macrophyte communities under light-reduction stress is how they balance carbon metabolism during photosynthesis and respiration. In turn, the dissolved organic carbon (DOC) released by these communities, which can be highly light-dependent, stands as a source of carbon, fuelling marine communities and playing an important role in the ocean carbon sequestration. This is the first study to evaluate light-reduction stress and recovery in the seagrass Zostera noltei and the macroalga Caulerpa prolifera. Light reduction led to a significant decrease in the production of both communities from autotrophic to heterotrophic. Results indicated that most of the DOC released by vegetated coastal communities comes from photosynthetic activity, and that the net DOC fluxes can be greatly affected by shading events. Finally, both communities showed resilience underpinned by high recovery but low resistance capacity, with C. prolifera showing the highest resilience to unfavourable light conditions., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Seagrass meadows, through their large capacity to sequester and store organic carbon in their sediments, contribute to mitigate climatic change. However, these ecosystems have experienced large losses and degradation worldwide due to anthropogenic and natural impacts and they are among the most threatened ecosystems on Earth. When a meadow is impacted, the vegetation is partial- or completely lost, and the sediment is exposed to the atmosphere or water column, resulting in the erosion and remineralisation of the carbon stored. This paper addresses the effects of the construction of coastal infrastructures on sediment properties, organic carbon, and total nitrogen stocks of intertidal seagrass meadows, as well as the size of such stocks in relation to meadow establishing time (recently and old established meadows). Three intertidal seagrass meadows impacted by coastal constructions (with 0% seagrass cover at present) and three adjacent non-impacted old-established meadows (with 100% seagrass cover at present) were studied along with an area of bare sediment and two recent-established seagrass meadows. We observed that the non-impacted areas presented 3-fold higher percentage of mud and 1.5 times higher sedimentary organic carbon stock than impacted areas. Although the impacted area was relatively small (0.05-0.07 ha), coastal infrastructures caused a significant reduction of the sedimentary carbon stock, between 1.1 and 2.2 Mg OC, and a total loss of the carbon sequestration capacity of the impacted meadow. We also found that the organic carbon stock and total nitrogen stock of the recent-established meadow were 30% lower than those of the old-established ones, indicating that OC and TN accumulation within the meadows is a continuous process, which has important consequences for conservation and restoration actions. These results contribute to understanding the spatial variability of blue carbon and nitrogen stocks in coastal systems highly impacted by urban development., Competing Interests: Declaration of competing interest Authors declare that there is no conflict of interest., (Copyright © 2022 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Seagrass beds act as blue carbon sinks globally as they enhance the trapping of recalcitrant (i.e., low biodegradability) organic carbon in their sediments. Recent studies also show that the recalcitrant fraction of the dissolved organic carbon (DOC) pool in seawater has an important role as long-term carbon sequestration in oceans. Although seagrasses are known for the large amount of DOC they export, little attention has been given to its biodegradability, which ultimately determinates its fate in the coastal carbon cycle. In turn, invasive algae are a major global concern in seagrass ecosystems since they can deeply modify their structure and functions, which may affect carbon metabolism and DOC release. This work assesses how the presence of Halimeda incrassata, an invasive tropical calcareous macroalga, modifies carbon metabolism and DOC fluxes in invaded areas dominated by the seagrass Cymodocea nodosa. Our results show that stands with the presence of this seagrass (i.e., both monospecific and mixed meadow) had the highest production values, acting as high DOC producers in both winter (mainly of labile DOC; DOC L ) and summer (mainly as recalcitrant DOC; DOC R ). In contrast, monospecific H. incrassata beds exhibited low production values, and the presence of this macroalga (either as monospecific beds or mixed with C. nodosa) triggered the shift from a net DOC-producing-system in summer (mainly DOC L ) to a net DOC-consuming-system in winter. This work thus suggests that C. nodosa meadows have the potential to export a significant fraction of both labile and recalcitrant DOC, and that the spread of this invasive alga might decrease the C export capacity of seagrass meadows. Such shift would imply the reduction of a quick and efficient transfer of carbon and energy to higher trophic levels, and might reduce the blue carbon potential of seagrasses as dissolved form in the water column., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 The Authors. Published by Elsevier B.V. All rights reserved.)

Cadiz Bay is a shallow mesotidal lagoon with extensive populations of the seagrass Cymodocea nodosa at intertidal and shallow subtidal elevations. This work aims to understand the mechanisms behind the resilience of this species to gradual sea level rise by studying its acclimation capacity to depth along the shallow littoral, and therefore, to gradual variations in the light environment. To address this objective, these populations have been monitored seasonally over a 10 year period, representing the longest seasonal database available in the literature for this species. The monitoring included populations at 0.4, -0.08 and -0.5 m LAT. The results show that C. nodosa has a strong seasonality for demographic and shoot dynamic properties - with longer shoots and larger growth in summer (high temperature) than in winter (low temperature), but also some losses. Moreover, shoots have different leaf morphometry depending on depth, with small and dense shoots in the intertidal areas (0.4 m) and sparse large shoots in the subtidal ones (-0.08 and 0.5 m). These differences in morphometry and shoot dynamic properties, combined with the differences in shoot density, explain the lack of differences in meadow production balance (i.e. meadow growth - meadow losses) between the intertidal (0.4 m) and the deepest population (-0.5 m), supporting the long term resilience of Cymodocea nodosa in Cadiz Bay. This study contributes to the understanding of the mechanisms behind seagrass stability and resilience, which is particularly important towards predicting the effects of climate change on these key coastal ecosystems, and also highlights the value of continuous long-term monitoring efforts to evaluate seagrass trajectories., (Copyright © 2021 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Seaweeds (macroalgae) are, together with microalgae, main contributors to the Earth's production of organic matter and atmospheric oxygen as well as fixation of carbon dioxide. In addition, they contain a bounty of fibres and minerals, as well as macro- and micronutrients that can serve both technical and medicinal purposes, as well as be a healthy and nutritious food for humans and animals. It is therefore natural that seaweeds and humans have had a myriad of interwoven relationships both on evolutionary timescales as well as in recent millennia and centuries all the way into the Anthropocene. It is no wonder that seaweeds have also entered and served as a saviour for humankind around the globe in many periods of severe needs and crises. Indeed, they have sometimes been the last resort, be it during times of famine, warfare, outbreak of diseases, nuclear accidents, or as components of securing the fabric of social stability. The present topical review presents testimony from the history of human interaction with seaweeds to the way humankind has, over and over again, been 'saved by seaweeds'. It remains a historical fact that in extreme conditions, such as shortage and wars, humans have turned to seaweeds in times of 'needs must' and created new opportunities for their uses in order to mitigate disasters. Lessons to be learned from this history can be used as reminders and inspiration, and as a guide as how to turn to seaweeds in current and inevitable, future times of crises, not least for the present needs of how to deal with changing climates and the pressing challenges of sustainable and healthy eating., Competing Interests: Conflict of interestThe authors declare that they have no conflict of interest., (© Springer Nature B.V. 2020.)

Seagrass meadows, key ecosystems supporting fisheries, carbon sequestration and coastal protection, are globally threatened. In Europe, loss and recovery of seagrasses are reported, but the changes in extent and density at the continental scale remain unclear. Here we collate assessments of changes from 1869 to 2016 and show that 1/3 of European seagrass area was lost due to disease, deteriorated water quality, and coastal development, with losses peaking in the 1970s and 1980s. Since then, loss rates slowed down for most of the species and fast-growing species recovered in some locations, making the net rate of change in seagrass area experience a reversal in the 2000s, while density metrics improved or remained stable in most sites. Our results demonstrate that decline is not the generalised state among seagrasses nowadays in Europe, in contrast with global assessments, and that deceleration and reversal of declining trends is possible, expectingly bringing back the services they provide.

Seagrass shoots interact with hydrodynamic forces and thereby a positively or negatively influence the survival of associated species. The modification of these forces indirectly alters the physical transport and flux of edible particles within seagrass meadows, which will influence the growth and survivorship of associated filter-feeding organisms. The present work contributes to gaining insight into the mechanisms controlling the availability of resources for filter feeders inhabiting seagrass canopies, both from physical (influenced by seagrass density and patchiness) and biological (regulated by filter feeder density) perspectives. A factorial experiment was conducted in a large racetrack flume, which combined changes in hydrodynamic conditions, chlorophyll a concentration in the water and food intake rate (FIR) in a model active filter-feeding organism (the cockle). Results showed that seagrass density and patchiness modified both hydrodynamic forces and availability of resources for filter feeders. Chlorophyll a water content decreased to 50% of the initial value when densities of both seagrass shoots and cockles were high. Also, filter feeder density controlled resource availability within seagrass patches, depending on its spatial position within the racetrack flume. Under high density of filter-feeding organisms, chlorophyll a levels were lower between patches. This suggests that the pumping activity of cockles (i.e. biomixing) is an emergent key factor affecting both resource availability and FIR for filter feeders in dense canopies. Applying our results to natural conditions, we suggest the existence of a direct correlation between habitat complexity (i.e. shoot density and degree of patchiness) and filter feeders density. Fragmented and low-density patches seem to offer both greater protection from hydrodynamic forces and higher resource availability. In denser patches, however, resources are allocated mostly within the canopy, which would benefit filter feeders if they occurred at low densities, but would be limiting when filter feeder were at high densities.

An in-depth knowledge of solutes advection and turbulent diffusion is crucial to estimate dispersion area and retention time (t(R)) of pollutants within seagrass habitats. However, there is little knowledge on the influence of seagrass habitat fragmentation on such mechanisms. A set of dye tracer experiments and acoustic Doppler velocimeter measurements (ADV) were conducted. Solute transport conditions were compared in between fragmented (FM) vs homogeneous (HM) intertidal meadows, and in vertical gradients (canopy vs overlaying flow). Results showed the highest horizontal diffusion coefficient (K(y), c.a. 10(-3)m(2)s(-1)) on FM and at the canopy-water column interface, whereas t(R) (2.6-5.6 min) was not affected by fragmentation. It suggests that (1) FM are more vulnerable to pollution events in terms of dispersion area and (2) at low tide, advection rather than turbulent diffusion determines t(R). Furthermore, Taylor's theorem is revealed as a powerful tool to analyze vertical gradients on K(y) within seagrass canopies., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

Disaggregating seagrass meadows and studying its components separately (clones, ramets, shoots) can provide us insights on meadow dynamics and growth patterns. The clonal growth, dependent upon clonal rules may regulate and impose constraints to plant architecture and, therefore, determine how individual clones evolve into the environment. In order to investigate the relationship between clonal growth rules and clone architecture, the belowground network architecture of single-clones of the seagrass Zostera noltii was studied. Networks were traced in situ after washing out the overlying sediment, and network characteristics were measured using digital analysis: area covered by clone, total rhizome length, type of rhizomatic axes (main, secondary, tertiary, quaternary), number and length of the internodes, branching angles and branching frequencies. This approach revealed that Z. noltii is able to develop into large clones integrating up to 300 internodes, 676 cm of rhizome, 208 shoots and 4,300 cm 2 of plant area. Internodal length depended on both, the distance to the apical shoot (time effect) and the axes type (apical dominance effect). However, average branching angle was independent of axis type (average 58.3 ± 0.75), but varied significantly depending on the distance from the apical shoot. This average branching angle allows Z. noltii maximize the rate of centrifugal expansion, maintaining a high density in colonized areas to produce close stands but also minimizing the investment in belowground biomass and ramets overlapping. The clonal architecture of Z. noltii seems to be regulated by the interaction of both, apical dominance strength and clonal integration distance. Moreover, clonal growth rules and growth pattern seem to constrain clonality through (clonal) plant architecture regulations (i.e. branching is restricted in secondary axes, similar average branching angles regardless the axes, the higher the distance to the apex the higher the number of internodes in secondary axes, shorter internodes in secondary and tertiary axes). Future research efforts should focus on how these complex relationships between apical dominance and clonal integration interact to elucidate the temporal (seasonal) and spatial scales of both processes and the outcome at the plant architectural level.

Abstract The extent of clonality within populations strongly influences their spatial genetic structure (SGS), yet this is hardly ever thoroughly analysed. We employed spatial autocorrelation analysis to study effects of sexual and clonal reproduction on dispersal of the dioecious seagrass Cymodocea nodosa. Analyses were performed both at genet level (i.e. excluding clonal repeats) and at ramet level. Clonal structure was characterized by the clonal subrange, a spatial measure of the linear limits where clonality still affects SGS. We show that the clonal subrange is equivalent to the distance where the probability of clonal identity approaches zero. This combined approach was applied to two meadows with different levels of disturbance, Cadiz (stable) and Alfacs (disturbed). Genotypic richness, the proportion of the sample representing distinct genotypes, was moderate (0.38 Cadiz, 0.46 Alfacs) mostly due to dominance of a few clones. Expected heterozygosities were comparable to those found in other clonal plants. SGS analyses at the genet level revealed extremely restricted gene dispersal in Cadiz (Sp = 0.052, a statistic reflecting the decrease of pairwise kinship with distance), the strongest SGS found for seagrass species, comparable only to values for selfing herbaceous land plants. At Cadiz the clonal subrange extended across shorter distances (20-25 m) than in Alfacs (30-35 m). Comparisons of sexual and vegetative components of gene dispersal suggest that, as a dispersal vector within meadows, clonal spread is at least as important as sexual reproduction. The restricted dispersal and SGS pattern in both meadows indicates that the species follows a repeated seedling recruitment strategy.

The effects of light reduction [100%, 25%, 10% and 1% mean daily-integrated photon irradiance (I 0 )] by Ulva rigida C. Agardh canopies on carbon balance, sugar-related enzymes and proteolytic activities of the seagrass Zostera noltii Hornem. were investigated. Shaded plants showed negative net growth and starch was mobilized in both above- and below-ground tissues. Sucrose declined in below-ground parts under severe light deprivation (10% and 1% I 0 ), but was accumulated in above-ground parts. Mobilization of the non-structural carbohydrates (sucrose and starch) was explained by changes in activities of sucrose synthase (SuSy, EC 2.4.1.13) and sucrose-phosphate synthase (SPS, EC 2.4.2.24). Under severe light reduction, the capacity of above-ground tissues for sucrose formation and export declined, indicated by the lowest SPS activity. In contrast, severe light reduction increased the 'sink strength' of below-ground tissues, demonstrated by the highest SuSy activities, and diminished the capacity for sucrose resynthesis from starch breakdown, as the lowest SPS activity was observed under low light. These results suggest a cessation of sucrose transport throughout the plant under extreme light limitation, the carbon supply being dependent on the starch breakdown in each tissue. The response of Z. noltii to gradual light reduction was co-ordinated at the whole-plant level, since an enhancement of proteolytic activities induced by carbon starvation in both above- and below-ground tissues was also recorded during prolonged light deprivation. Therefore, carbon mobilization was accompanied by enhanced protein turnover and changes in metabolic pathways.

Within the sheltered creeks of Cádiz bay, Ulva thalli form extended mat-like canopies. The effect of solar ultraviolet radiation on photosynthetic activity, the composition of photosynthetic and xanthophyll cycle pigments, and the amount of RubisCO, chaperonin 60 (CPN 60), and the induction of DNA damage in Ulva aff. rotundata Bliding from southern Spain was assessed in the field. Samples collected from the natural community were covered by screening filters, generating different radiation conditions. During daily cycles, individual thalli showed photoinhibitory effects of the natural solar radiation. This inhibition was even more pronounced in samples only exposed to photosynthetically active radiation (PAR). Strongly increased heat dissipation in these samples indicated the activity of regulatory mechanisms involved in dynamic photoinhibition. Adverse effects of UV-B radiation on photosynthesis were only observed in combination with high levels of PAR, indicating the synergistic effects of the two wavelength ranges. In samples exposed either to PAR+UV-A or to UV-B+UV-A without PAR, no inhibition of photosynthetic quantum yield was found in the course of the day. At the natural site, the top layer of the mat-like canopies is generally completely bleached. Artificially designed Ulva canopies exhibited fast bleaching of the top layer under the natural solar radiation conditions, while this was not observed in canopies either shielded from UV or from PAR. The bleached first layer of the canopies acts as a selective UV-B filter, and thus prevents subcanopy thalli from exposure to harmful radiation. This was confirmed by the differences in photosynthetic activity, pigment composition, and the concentration of RubisCO in thalli with different positions within the canopy. In addition, the induction of the stress protein CPN 60 under UV exposure and the low accumulation of DNA damage indicate the presence of physiological protection mechanisms against harmful UV-B. A mechanism of UV-B-induced inhibition of photosynthesis under field conditions is proposed.