A trait-based framework for seagrass ecology: Trends and prospects.

In the last three decades, quantitative approaches that rely on organism traits instead of taxonomy have advanced different fields of ecological research through establishing the mechanistic links between environmental drivers, functional traits, and ecosystem functions. A research subfield where trait-based approaches have been frequently used but poorly synthesized is the ecology of seagrasses; marine angiosperms that colonized the ocean 100M YA and today make up productive yet threatened coastal ecosystems globally. Here, we compiled a comprehensive trait-based response-effect framework (TBF) which builds on previous concepts and ideas, including the use of traits for the study of community assembly processes, from dispersal and response to abiotic and biotic factors, to ecosystem function and service provision. We then apply this framework to the global seagrass literature, using a systematic review to identify the strengths, gaps, and opportunities of the field. Seagrass trait research has mostly focused on the effect of environmental drivers on traits, i.e., "environmental filtering" (72%), whereas links between traits and functions are less common (26.9%). Despite the richness of trait-based data available, concepts related to TBFs are rare in the seagrass literature (15% of studies), including the relative importance of neutral and niche assembly processes, or the influence of trait dominance or complementarity in ecosystem function provision. These knowledge gaps indicate ample potential for further research, highlighting the need to understand the links between the unique traits of seagrasses and the ecosystem services they provide.

Methane emissions from macrophyte beach wrack on Baltic seashores.

Beach wrack of marine macrophytes is a natural component of many beaches. To test if such wrack emits the potent greenhouse gas methane, field measurements were made at different seasons on beach wrack depositions of different ages, exposure, and distance from the water. Methane emissions varied greatly, from 0 to 176 mg CH4-C m-2 day-1, with a clear positive correlation between emission and temperature. Dry wrack had lower emissions than wet. Using temperature data from 2016 to 2020, seasonal changes in fluxes were calculated for a natural wrack accumulation area. Such calculated average emissions were close to zero during winter, but peaked in summer, with very high emissions when daily temperatures exceeded 20 °C. We conclude that waterlogged beach wrack significantly contributes to greenhouse gas emissions and that emissions might drastically increase with increasing global temperatures. When beach wrack is collected into heaps away from the water, the emissions are however close to zero.

Impacts of land-use change and urban development on carbon sequestration in tropical seagrass meadow sediments.

Seagrass meadows store significant carbon stocks at a global scale, but land-use change and other anthropogenic activities can alter the natural process of organic carbon (Corg) accumulation. Here, we assessed the carbon accumulation history of two seagrass meadows in Zanzibar (Tanzania) that have experienced different degrees of disturbance. The meadow at Stone Town has been highly exposed to urban development during the 20th century, while the Mbweni meadow is located in an area with relatively low impacts but historical clearing of adjacent mangroves. The results showed that the two sites had similar sedimentary Corg accumulation rates (22-25 g m-2 yr-1) since the 1940s, while during the last two decades (∼1998 until 2018) they exhibited 24-30% higher accumulation of Corg, which was linked to shifts in Corg sources. The increase in the δ13C isotopic signature of sedimentary Corg (towards a higher seagrass contribution) at the Stone Town site since 1998 points to improved seagrass meadow conditions and Corg accumulation capacity of the meadow after the relocation of a major sewage outlet in the mid-1990s. In contrast, the decrease in the δ13C signatures of sedimentary Corg in the Mbweni meadow since the early 2010s was likely linked to increased Corg run-off of mangrove/terrestrial material following mangrove deforestation. This study exemplifies two different pathways by which land-based human activities can alter the carbon storage capacity of seagrass meadows (i.e. sewage waste management and mangrove deforestation) and showcases opportunities for management of vegetated coastal Corg sinks.

Seagrass meadows mixed with calcareous algae have higher plant productivity and sedimentary blue carbon storage.

Seagrass meadows capture and store large amounts of carbon in the sediment beneath, thereby serving as efficient sinks of atmospheric CO2. Carbon sequestration levels may however differ greatly among meadows depending on, among other factors, the plant community composition. Tropical seagrass meadows are often intermixed with macroalgae, many of which are calcareous, which may compete with seagrass for nutrients, light, and space. While the photosynthetic CO2 uptake by both seagrasses and calcareous algae may increase the overall calcification in the system (by increasing the calcium carbonate saturation state, Ω), the calcification process of calcareous algae may lead to a release of CO2, thereby affecting both productivity and calcification, and eventually also the meadows' carbon storage. This study estimated how plant productivity, CaCO3 production, and sediment carbon levels were affected by plant community composition (seagrass and calcareous algae) in a tropical seagrass-dominated embayment (Zanzibar, Tanzania). Overall, the patterns of variability in productivity differed between the plant types, with net areal biomass productivity being highest in meadows containing both seagrass and calcareous algae. Low and moderate densities of calcareous algae enhanced seagrass biomass growth, while the presence of seagrass reduced the productivity of calcareous algae but increased their CaCO3 content. Sedimentary carbon levels were highest when seagrasses were mixed with low or moderate cover of calcareous algae. The findings show that plant community composition can be an important driver for ecosystem productivity and blue carbon sequestration.

The amount of light reaching the leaves in seagrass (Zostera marina) meadows.

Seagrass meadows, and other submerged vegetated habitats, support a wide range of essential ecological services, but the true extents of these services are in many ways still not quantified. One important tool needed to assess and model many of these services is accurate estimations of the systems´ primary productivity. Such productivity estimations require an understanding of the underwater light field, especially regarding the amount of light that actually reaches the plants' photosynthetic tissue. In this study, we tested a simple practical approach to estimate leaf light exposure, relative to incoming light at the canopy, by attaching light sensitive film at different positions on leaves of Zostera marina, eelgrass, in four seagrass meadows composed of different shoot density and at two different depths. We found that the light reaching the leaves decreased linearly down through the canopy. While the upper parts of the leaves received approximately the same level of light (photosynthetic photon flux density, PPFD) as recorded with a PAR meter at the canopy top, the average light that the seagrass leaves were exposed to varied between 40 and 60% of the light on top of the canopy, with an overall average of 48%. We recommend that actual light interception is measured when assessing or modelling light depending processes in submerged vegetation, but if this is not achievable a rough estimation for vegetation similar to Z. marina would be to use a correction factor of 0.5 to compensate for the reduced light due to leaf orientation and internal shading.

A temporal record of microplastic pollution in Mediterranean seagrass soils.

Plastic pollution is emerging as a potential threat to the marine environment. In the current study, we selected seagrass meadows, known to efficiently trap organic and inorganic particles, to investigate the concentrations and dynamics of microplastics in their soil. We assessed microplastic contamination and accumulation in 210Pb dated soil cores collected in Posidonia oceanica meadows at three locations along the Spanish Mediterranean coast, with two sites located in the Almería region (Agua Amarga and Roquetas) and one at Cabrera Island (Santa Maria). Almería is known for its intense agricultural industry with 30 000 ha of plastic-covered greenhouses, while the Cabrera Island is situated far from urban areas. Microplastics were extracted using enzymatic digestion and density separation. The particles were characterized by visual identification and with Fourier-transformed infrared (FTIR) spectroscopy, and related to soil age-depth chronologies. Our findings showed that the microplastic contamination and accumulation was negligible until the mid-1970s, after which plastic particles increased dramatically, with the highest concentrations of microplastic particles (MPP) found in the recent (since 2012) surface soil of Agua Amarga (3819 MPP kg-1), followed by the top-most layers of the soil of the meadows in Roquetas (2173 kg-1) and Santa Maria (68-362 kg-1). The highest accumulation rate was seen in the Roquetas site (8832 MPP m-2 yr-1). The increase in microplastics in the seagrass soil was associated to land-use change following the intensification of the agricultural industry in the area, with a clear relationship between the development of the greenhouse industry in Almería and the concentration of microplastics in the historical soil record. This study shows a direct linkage between intense anthropogenic activity, an extensive use of plastics and high plastic contamination in coastal marine ecosystems such as seagrass meadows. We highlight the need of proper waste management to protect the coastal environment from continuous pollution.

The influence of hydrodynamic exposure on carbon storage and nutrient retention in eelgrass (Zostera marina L.) meadows on the Swedish Skagerrak coast.

Cold-temperate seagrass (Zostera marina) meadows provide several important ecosystem services, including trapping and storage of sedimentary organic carbon and nutrients. However, seagrass meadows are rapidly decreasing worldwide and there is a pressing need for protective management of the meadows and the organic matter sinks they create. Their carbon and nutrient storage potential must be properly evaluated, both at present situation and under future climate change impacts. In this study, we assessed the effect of wave exposure on sedimentary carbon and nitrogen accumulation using existing data from 53 Z. marina meadows at the Swedish west coast. We found that meadows with higher hydrodynamic exposure had larger absolute organic carbon and nitrogen stocks (at 0-25 cm depth). This can be explained by a hydrodynamically induced sediment compaction in more exposed sites, resulting in increased sediment density and higher accumulation (per unit volume) of sedimentary organic carbon and nitrogen. With higher sediment density, the erosion threshold is assumed to increase, and as climate change-induced storms are predicted to be more common, we suggest that wave exposed meadows can be more resilient toward storms and might therefore be even more important as carbon- and nutrient sinks in the future.

An experimental assessment of algal calcification as a potential source of atmospheric CO2.

Marine vegetated ecosystems such as seagrass meadows are increasingly acknowledged as important carbon sinks based on their ability to capture and store atmospheric carbon dioxide, thereby contributing to climate change mitigation. Most studies on carbon storage in marine ecosystems have focused on organic carbon, leaving inorganic carbon processes such as calcification unaccounted for, despite of their critical role in the global carbon budget. This is probably because of uncertainties regarding the role of calcification in marine carbon budgets as either atmospheric CO2 source or sink. Here, we conducted a laboratory experiment to investigate the influence of a calcifying alga (Corallina officinalis L.) on seawater carbon content, using a non-calcifying alga (Ulva lactuca L.) as a control. In a first part, algae were incubated separately while measuring changes in seawater pH, total alkalinity (TA) and total dissolved inorganic carbon (DIC). The amount of carbon used in photosynthetic uptake and production of CaCO3 was then calculated. In a second, directly following, part the algae were removed and DIC levels were allowed to equilibrate with air until the pH stabilized and the loss of CO2 to air was calculated as the difference in total DIC from the start of part one, to the end of the second part. The results showed that C. officinalis caused a significant and persistent reduction in total dissolved inorganic carbon (DIC), TA and seawater pH, while no such permanent changes were caused by U. lactuca. These findings indicate that calcification can release a significant amount of CO2 to the atmosphere and thereby possibly counteract the carbon sequestration in marine vegetated ecosystems if this CO2 is not re-fixed in the system. Our research emphasises the importance of considering algal calcification in future assessments on carbon storage in coastal areas.

Methane emission and sulfide levels increase in tropical seagrass sediments during temperature stress: A mesocosm experiment.

Climate change-induced ocean warming is expected to greatly affect carbon dynamics and sequestration in vegetated shallow waters, especially in the upper subtidal where water temperatures may fluctuate considerably and can reach high levels at low tides. This might alter the greenhouse gas balance and significantly reduce the carbon sink potential of tropical seagrass meadows. In order to assess such consequences, we simulated temperature stress during low tide exposures by subjecting seagrass plants (Thalassia hemprichii) and associated sediments to elevated midday temperature spikes (31, 35, 37, 40, and 45°C) for seven consecutive days in an outdoor mesocosm setup. During the experiment, methane release from the sediment surface was estimated using gas chromatography. Sulfide concentration in the sediment pore water was determined spectrophotometrically, and the plant's photosynthetic capacity as electron transport rate (ETR), and maximum quantum yield (Fv/Fm) was assessed using pulse amplitude modulated (PAM) fluorometry. The highest temperature treatments (40 and 45°C) had a clear positive effect on methane emission and the level of sulfide in the sediment and, at the same time, clear negative effects on the photosynthetic performance of seagrass plants. The effects observed by temperature stress were immediate (within hours) and seen in all response variables, including ETR, Fv/Fm, methane emission, and sulfide levels. In addition, both the methane emission and the size of the sulfide pool were already negatively correlated with changes in the photosynthetic rate (ETR) during the first day, and with time, the correlations became stronger. These findings show that increased temperature will reduce primary productivity and increase methane and sulfide levels. Future increases in the frequency and severity of extreme temperature events could hence reduce the climate mitigation capacity of tropical seagrass meadows by reducing CO2 sequestration, increase damage from sulfide toxicity, and induce the release of larger amounts of methane.

Above-below surface interactions mediate effects of seagrass disturbance on meiobenthic diversity, nematode and polychaete trophic structure.

Ecological interactions between aquatic plants and sediment communities can shape the structure and function of natural systems. Currently, we do not fully understand how seagrass habitat degradation impacts the biodiversity of belowground sediment communities. Here, we evaluated indirect effects of disturbance of seagrass meadows on meiobenthic community composition, with a five-month in situ experiment in a tropical seagrass meadow. Disturbance was created by reducing light availability (two levels of shading), and by mimicking grazing events (two levels) to assess impacts on meiobenthic diversity using high-throughput sequencing of 18S rRNA amplicons. Both shading and simulated grazing had an effect on meiobenthic community structure, mediated by seagrass-associated biotic drivers and sediment abiotic variables. Additionally, shading substantially altered the trophic structure of the nematode community. Our findings show that degradation of seagrass meadows can alter benthic community structure in coastal areas with potential impacts to ecosystem functions mediated by meiobenthos in marine sediments.

Estimation of a whole plant Q10 to assess seagrass productivity during temperature shifts.

Through respiration and photosynthesis, seagrass meadows contribute greatly to carbon and oxygen fluxes in shallow coastal waters. There is increasing concern about how shallow-water primary producers will react to a near-future climate scenario with increased temperature variation. When modelling primary productivity under high temperature variability, Q10 values are commonly used to predict rate changes depending on biophysical factors. Q10 values are often assumed to be constant and around 2.0 (i.e. a doubling of the rate with a temperature increase of 10 °C). We aimed to establish how the gas exchange of seagrass (Zostera marina) tissues at various maturity stages would respond over a broad range of temperatures. Seagrass shoot maturity stage clearly affected respiration and apparent photosynthesis, and the Q10 results indicated a skewed balance between the two processes, with a higher photosynthetic Q10 during periods of elevated temperatures. When estimating whole-plant Q10 in a realistic maximal temperature range, we found that the overall response of a seagrass plant's net O2 exchange balance can be as much as three to four times higher than under ambient temperatures. Our findings indicate that plant tissue age and temperature should be considered when assessing and modelling carbon and oxygen fluctuations in vegetated coastal areas.

High midday temperature stress has stronger effects on biomass than on photosynthesis: A mesocosm experiment on four tropical seagrass species.

The effect of repeated midday temperature stress on the photosynthetic performance and biomass production of seagrass was studied in a mesocosm setup with four common tropical species, including Thalassia hemprichii, Cymodocea serrulata, Enhalus acoroides, and Thalassodendron ciliatum. To mimic natural conditions during low tides, the plants were exposed to temperature spikes of different maximal temperatures, that is, ambient (29-33°C), 34, 36, 40, and 45°C, during three midday hours for seven consecutive days. At temperatures of up to 36°C, all species could maintain full photosynthetic rates (measured as the electron transport rate, ETR) throughout the experiment without displaying any obvious photosynthetic stress responses (measured as declining maximal quantum yield, Fv/Fm). All species except T. ciliatum could also withstand 40°C, and only at 45°C did all species display significantly lower photosynthetic rates and declining Fv/Fm. Biomass estimation, however, revealed a different pattern, where significant losses of both above- and belowground seagrass biomass occurred in all species at both 40 and 45°C (except for C. serrulata in the 40°C treatment). Biomass losses were clearly higher in the shoots than in the belowground root-rhizome complex. The findings indicate that, although tropical seagrasses presently can cope with high midday temperature stress, a few degrees increase in maximum daily temperature could cause significant losses in seagrass biomass and productivity.

Shading and simulated grazing increase the sulphide pool and methane emission in a tropical seagrass meadow.

Though seagrass meadows are among the most productive habitats in the world, contributing substantially to long-term carbon storage, studies of the effects of critical disturbances on the fate of carbon sequestered in the sediment and biomass of these meadows are scarce. In a manipulative in situ experiment, we studied the effects of successive loss of seagrass biomass as a result of shading and simulated grazing at two intensity levels on sulphide (H2S) content and methane (CH4) emission in a tropical seagrass meadow in Zanzibar (Tanzania). In all disturbed treatments, we found a several-fold increase in both the sulphide concentration of the sediment pore-water and the methane emissions from the sediment surface (except for CH4 emissions in the low-shading treatment). This could be due to the ongoing degradation of belowground biomass shed by the seagrass plants, supporting the production of both sulphate-reducing bacteria and methanogens, possibly exacerbated by the loss of downwards oxygen transport via seagrass plants. The worldwide rapid loss of seagrass areas due to anthropogenic activities may therefore have significant effects on carbon sink-source relationships within coastal seas.

Contribution of seagrass plants to CO2 capture in a tropical seagrass meadow under experimental disturbance.

Coastal vegetative habitats are known to be highly productive environments with a high ability to capture and store carbon. During disturbance this important function could be compromised as plant photosynthetic capacity, biomass, and/or growth are reduced. To evaluate effects of disturbance on CO2 capture in plants we performed a five-month manipulative experiment in a tropical seagrass (Thalassia hemprichii) meadow exposed to two intensity levels of shading and simulated grazing. We assessed CO2 capture potential (as net CO2 fixation) using areal productivity calculated from continuous measurements of diel photosynthetic rates, and estimates of plant morphology, biomass and productivity/respiration (P/R) ratios (from the literature). To better understand the plant capacity to coping with level of disturbance we also measured plant growth and resource allocation. We observed substantial reductions in seagrass areal productivity, biomass, and leaf area that together resulted in a negative daily carbon balance in the two shading treatments as well as in the high-intensity simulated grazing treatment. Additionally, based on the concentrations of soluble carbohydrates and starch in the rhizomes, we found that the main reserve sources for plant growth were reduced in all treatments except for the low-intensity simulated grazing treatment. If permanent, these combined adverse effects will reduce the plants' resilience and capacity to recover after disturbance. This might in turn have long-lasting and devastating effects on important ecosystem functions, including the carbon sequestration capacity of the seagrass system.

Respiratory oxygen consumption in the seagrass Zostera marina varies on a diel basis and is partly affected by light.

The seagrass Zostera marina is an important marine ecosystem engineer, greatly influencing oxygen and carbon fluctuations in temperate coastal areas. Although photosynthetically driven gas fluxes are well studied, the impact of the plant's mitochondrial respiration on overall CO2 and O2 fluxes in marine vegetated areas is not yet understood. Likewise, the gene expression in relation to the respiratory pathway has not been well analyzed in seagrasses. This study uses a combined approach, studying respiratory oxygen consumption rates in darkness simultaneously with changes in gene expression, with the aim of examining how respiratory oxygen consumption fluctuates on a diel basis. Measurements were first made in a field study where samples were taken directly from the ocean to the laboratory for estimations of respiratory rates. This was followed by a laboratory study where measurements of respiration and expression of genes known to be involved in mitochondrial respiration were conducted for 5 days under light conditions mimicking natural summer light (i.e., 15 h of light and 9 h of darkness), followed by 3 days of constant darkness to detect the presence of a potential circadian clock. In the field study, there was a clear diel variation in respiratory oxygen consumption with the highest rates in the late evening and at night (0.766 and 0.869 µmol O2 m-2 s-1, respectively). These repetitive diel patterns were not seen in the laboratory, where water conditions (temperature, pH, and oxygen) showed minor fluctuations and only light varied. The gene expression analysis did not give clear evidence on drivers behind the respiratory fluxes; however, expression levels of the selected genes generally increased when the seagrass was kept in constant darkness. While light may influence mitochondrial respiratory fluxes, it appears that other environmental factors (e.g., temperature, pH, or oxygen) could be of significance too. As seagrasses substantially alter the proportions of both oxygen and inorganic carbon in the water column and respiration is a great driver of these alterations, we propose that acknowledging the presence of respiratory fluctuations in nature should be considered when estimating coastal carbon budgets. As dark respiration in field at midnight was approximately doubled from that of midday, great over-, or underestimations of the respiratory carbon dioxide release from seagrasses could be made if values are just obtained at one specific time point and considered constant.

Depth-specific fluctuations of gene expression and protein abundance modulate the photophysiology in the seagrass Posidonia oceanica.

Here we present the results of a multiple organizational level analysis conceived to identify acclimative/adaptive strategies exhibited by the seagrass Posidonia oceanica to the daily fluctuations in the light environment, at contrasting depths. We assessed changes in photophysiological parameters, leaf respiration, pigments, and protein and mRNA expression levels. The results show that the diel oscillations of P. oceanica photophysiological and respiratory responses were related to transcripts and proteins expression of the genes involved in those processes and that there was a response asynchrony between shallow and deep plants probably caused by the strong differences in the light environment. The photochemical pathway of energy use was more effective in shallow plants due to higher light availability, but these plants needed more investment in photoprotection and photorepair, requiring higher translation and protein synthesis than deep plants. The genetic differentiation between deep and shallow stands suggests the existence of locally adapted genotypes to contrasting light environments. The depth-specific diel rhythms of photosynthetic and respiratory processes, from molecular to physiological levels, must be considered in the management and conservation of these key coastal ecosystems.

Sediment Properties as Important Predictors of Carbon Storage in Zostera marina Meadows: A Comparison of Four European Areas.

Seagrass ecosystems are important natural carbon sinks but their efficiency varies greatly depending on species composition and environmental conditions. What causes this variation is not fully known and could have important implications for management and protection of the seagrass habitat to continue to act as a natural carbon sink. Here, we assessed sedimentary organic carbon in Zostera marina meadows (and adjacent unvegetated sediment) in four distinct areas of Europe (Gullmar Fjord on the Swedish Skagerrak coast, Askö in the Baltic Sea, Sozopol in the Black Sea and Ria Formosa in southern Portugal) down to ~35 cm depth. We also tested how sedimentary organic carbon in Z. marina meadows relates to different sediment characteristics, a range of seagrass-associated variables and water depth. The seagrass carbon storage varied greatly among areas, with an average organic carbon content ranging from 2.79 ± 0.50% in the Gullmar Fjord to 0.17 ± 0.02% in the area of Sozopol. We found that a high proportion of fine grain size, high porosity and low density of the sediment is strongly related to high carbon content in Z. marina sediment. We suggest that sediment properties should be included as an important factor when evaluating high priority areas in management of Z. marina generated carbon sinks.

The role of O2 as an electron acceptor alternative to CO2 in photosynthesis of the common marine angiosperm Zostera marina L.

This study investigates the role of O2 as an electron acceptor alternative to CO2 in photosynthesis of the common marine angiosperm Zostera marina L. Electron transport rates (ETRs) and non-photochemical quenching (NPQ) of Z. marina were measured under saturating irradiance in synthetic seawater containing 2.2 mM DIC and no DIC with different O2 levels (air-equilibrated levels, 3 % of air equilibrium and restored air-equilibrated levels). Lowering O2 did not affect ETR when DIC was provided, while it caused a decrease in ETR and an increase in NPQ in DIC-free media, indicating that O2 acted as an alternative electron acceptor under low DIC. The ETR and NPQ as a function of irradiance were subsequently assessed in synthetic seawater containing (1) 2.2 mM DIC, air-equilibrated O2; (2) saturating CO2, no O2; and (3) no DIC, air-equilibrated O2. These treatments were combined with glycolaldehyde pre-incubation. Glycolaldehyde caused a marked decrease in ETR in DIC-free medium, indicating significant electron flow supported by photorespiration. Combining glycolaldehyde with O2 depletion completely suppressed ETR suggesting the operation of the Mehler reaction, a possibility supported by the photosynthesis-dependent superoxide production. However, no notable effect of suppressing the Mehler reaction on NPQ was observed. It is concluded that during DIC-limiting conditions, such as those frequently occurring in the habitats of Z. marina, captured light energy exceeds what is utilised for the assimilation of available carbon, and photorespiration is a major alternative electron acceptor, while the contribution of the Mehler reaction is minor.

Photorespiration and carbon limitation determine productivity in temperate seagrasses.

The gross primary productivity of two seagrasses, Zostera marina and Ruppia maritima, and one green macroalga, Ulva intestinalis, was assessed in laboratory and field experiments to determine whether the photorespiratory pathway operates at a substantial level in these macrophytes and to what extent it is enhanced by naturally occurring shifts in dissolved inorganic carbon (DIC) and O2 in dense vegetation. To achieve these conditions in laboratory experiments, seawater was incubated with U. intestinalis in light to obtain a range of higher pH and O2 levels and lower DIC levels. Gross photosynthetic O2 evolution was then measured in this pretreated seawater (pH, 7.8-9.8; high to low DIC:O2 ratio) at both natural and low O2 concentrations (adjusted by N2 bubbling). The presence of photorespiration was indicated by a lower gross O2 evolution rate under natural O2 conditions than when O2 was reduced. In all three macrophytes, gross photosynthetic rates were negatively affected by higher pH and lower DIC. However, while both seagrasses exhibited significant photorespiratory activity at increasing pH values, the macroalga U. intestinalis exhibited no such activity. Rates of seagrass photosynthesis were then assessed in seawater collected from the natural habitats (i.e., shallow bays characterized by high macrophyte cover and by low DIC and high pH during daytime) and compared with open baymouth water conditions (where seawater DIC is in equilibrium with air, normal DIC, and pH). The gross photosynthetic rates of both seagrasses were significantly higher when incubated in the baymouth water, indicating that these grasses can be significantly carbon limited in shallow bays. Photorespiration was also detected in both seagrasses under shallow bay water conditions. Our findings indicate that natural carbon limitations caused by high community photosynthesis can enhance photorespiration and cause a significant decline in seagrass primary production in shallow waters.

Effects of CO(2) enrichment on photosynthesis, growth, and nitrogen metabolism of the seagrass Zostera noltii.

Seagrass ecosystems are expected to benefit from the global increase in CO(2) in the ocean because the photosynthetic rate of these plants may be C(i)-limited at the current CO(2) level. As well, it is expected that lower external pH will facilitate the nitrate uptake of seagrasses if nitrate is cotransported with H(+) across the membrane as in terrestrial plants. Here, we investigate the effects of CO(2) enrichment on both carbon and nitrogen metabolism of the seagrass Zostera noltii in a mesocosm experiment where plants were exposed for 5 months to two experimental CO(2) concentrations (360 and 700 ppm). Both the maximum photosynthetic rate (P(m)) and photosynthetic efficiency (α) were higher (1.3- and 4.1-fold, respectively) in plants exposed to CO(2)-enriched conditions. On the other hand, no significant effects of CO(2) enrichment on leaf growth rates were observed, probably due to nitrogen limitation as revealed by the low nitrogen content of leaves. The leaf ammonium uptake rate and glutamine synthetase activity were not significantly affected by increased CO(2) concentrations. On the other hand, the leaf nitrate uptake rate of plants exposed to CO(2)-enriched conditions was fourfold lower than the uptake of plants exposed to current CO(2) level, suggesting that in the seagrass Z. noltii nitrate is not cotransported with H(+) as in terrestrial plants. In contrast, the activity of nitrate reductase was threefold higher in plant leaves grown at high-CO(2) concentrations. Our results suggest that the global effects of CO(2) on seagrass production may be spatially heterogeneous and depend on the specific nitrogen availability of each system. Under a CO(2) increase scenario, the natural levels of nutrients will probably become limiting for Z. noltii. This potential limitation becomes more relevant because the expected positive effect of CO(2) increase on nitrate uptake rate was not confirmed.