Microbial diversity and keystone species drive soil nutrient cycling and multifunctionality following mangrove restoration.

Vegetation restoration exerts transformative effects on nutrient cycling, microbial communities, and ecosystem functions. While extensive research has been conducted on the significance of mangroves and their restoration efforts, the effectiveness of mangrove restoration in enhancing soil multifunctionality in degraded coastal wetlands remains unclear. Herein, we carried out a field experiment to explore the impacts of mangrove restoration and its chronosequence on soil microbial communities, keystone species, and soil multifunctionality, using unrestored aquaculture ponds as controls. The results revealed that mangrove restoration enhanced soil multifunctionality, with these positive effects progressively amplifying over the restoration chronosequence. Furthermore, mangrove restoration led to a substantial increase in microbial diversity and a reshaping of microbial community composition, increasing the relative abundance of dominant phyla such as Nitrospirae, Deferribacteres, and Fusobacteria. Soil multifunctionality exhibited positive correlations with microbial diversity, suggesting a link between variations in microbial diversity and soil multifunctionality. Metagenomic screening demonstrated that mangrove restoration resulted in a simultaneous increase in the abundance of nitrogen (N) related genes, such as N fixation (nirD/H/K), nitrification (pmoA-amoA/B/C), and denitrification (nirK, norB/C, narG/H, napA/B), as well as phosphorus (P)-related genes, including organic P mineralization (phnX/W, phoA/D/G, phnJ/N/P), inorganic P solubilization (gcd, ppx-gppA), and transporters (phnC/D/E, pstA/B/C/S)). The relationship between the abundance of keystone species (such as phnC/D/E) and restoration-induced changes in soil multifunctionality indicates that mangrove restoration enhances soil multifunctionality through an increase in the abundance of keystone species associated with N and P cycles. Additionally, it was observed that changes in microbial community and multifunctionality were largely associated with shifts in soil salinity. These findings demonstrate that mangrove restoration positively influences soil multifunctionality and shapes nutrient dynamics, microbial communities, and overall ecosystem resilience. As global efforts continue to focus on ecosystem restoration, understanding the complexity of mangrove-soil interactions is critical for effective nutrient management and mangrove conservation.

Moderate salinity improves the availability of soil P by regulating P-cycling microbial communities in coastal wetlands.

Accelerated sea-level rise is expected to cause the salinization of freshwater wetlands, but the responses to salinity of the availability of soil phosphorus (P) and of microbial genes involved in the cycling of P remain unexplored. We conducted a field experiment to investigate the effects of salinity on P cycling by soil microbial communities and their regulatory roles on P availability in coastal freshwater and brackish wetlands. Salinity was positively correlated with P availability, with higher concentrations of labile P but lower concentrations of moderately labile P in the brackish wetland. The diversity and richness of microbial communities involved in P cycling were higher in the brackish wetland than the freshwater wetland. Salinity substantially altered the composition of the P-cycling microbial community, in which those of the brackish wetland were separated from those of the freshwater wetland. Metagenomic sequence analysis indicated that functional genes involved in the solubilization of inorganic P and the subsequent transport and regulation of P were more abundant in coastal soils. The relative abundances of most of the target genes differed between the wetlands, with higher abundances of P-solubilization (gcd and ppa) and -mineralization (phoD, phy, and ugpQ) genes and lower abundances of P-transport genes (pstB, ugpA, ugpB, ugpE, and pit) in the brackish wetland. A significant positive correlation between the concentration of labile P and the abundances of the target genes suggested that salinity may, at least in part, improve P availability by regulating the P-cycling microbial community. Our results suggest that the P-cycling microbial community abundance and P availability respond positively to moderate increases in salinity by promoting the microbial solubilization and mineralization of soil P. Changes in microbial communities and microbially mediated P cycling may represent microbial strategies to adapt to moderate salinity levels, which in turn control soil function and nutrient balance.

Coastal degradation regulates the availability and diffusion kinetics of phosphorus at the sediment-water interface: Mechanisms and environmental implications.

Coastal wetlands have experienced considerable loss and degradation globally. However, how coastal degradation regulates sediment phosphorus (P) transformation and its underlying mechanisms remain largely unknown in subtropical coastal ecosystems. This study conducted seasonal field measurements using high-resolution diffusive gradient in thin films (DGT) and dialysis (Peeper) techniques, as well as a DGT-induced fluxes in sediments (DIFS) model, to evaluate the mobilization and diffusion of P along a degradation gradient ranging from pristine wetlands to moderately and severely degraded sites. We observed that sediment P is diminished by coastal degradation, and severely degraded sites exhibit a decline in the concentration of available P, despite the presence of distinct seasonal patterns. High-resolution data based on DGT/Peeper analysis revealed that labile P and soluble reactive P (SRP) concentrations varied from 0.0006 mg L-1 to 0.084 mg L-1 (mean 0.0147 mg L-1) and from 0.0128 mg L-1 to 0.1677 mg L-1 (mean 0.0536 mg L-1), respectively. Coastal degradation had a substantial impact on increasing SRP and labile P concentrations, particularly at severely degraded sites. Although severely degraded wetlands appeared to be P sinks (negative P flux at these sites), we did also observe positive diffusive flux in October, indicating that coastal degradation may accelerate the diffusion and remobilization of sediment P into overlying water. The simulations of the DIFS model provided compelling proof of the high resupply capacity of sediment P at severely degraded sites, as supported by the increased R and k-1 values but decreased Tc values. Taken together, these results suggest coastal degradation reduces the sediment P pool, primarily attributed to the strong remobilization of P from the sediment to porewater and overlying water by enhancing the resupply capability and diffusion kinetics. This acceleration induces nutrient loss which adversely impacts the water quality of the surrounding ecosystem. To reduce the adverse effects of coastal degradation, it is essential to adopt a combination of conservation, restoration, and management efforts designed to mitigate the risk of internal P loading and release, and ultimately maintain a regional nutrient balance.

Coastal conversion alters topsoil carbon, nitrogen and phosphorus stocks and stoichiometric balances in subtropical coastal wetlands.

The extensive conversion of coastal wetlands into agricultural and aquaculture areas has significant repercussions on soil nutrient balance. However, how coastal conversion specifically influences the dynamics and stoichiometry of topsoil carbon (C), nitrogen (N), and phosphorus (P) remains limited due to the considerable spatial variability and a lack of comprehensive field data. Here, we investigated the concentration and distribution of total C (TC), N (TN) and P (TP), along with their stoichiometric balance in four distinct coastal landscapes, including natural marshes and tidal flats, as well as converted agricultural croplands and ponds. The results revealed that converted croplands and ponds exhibited significantly higher concentrations of soil C, N and P, particularly in comparison to tidal flats. Furthermore, croplands and ponds have higher topsoil C stocks than tidal flats, but little difference or even lose stored C compared to marshes. Cropland soils showed considerably higher levels of available N (NH4+-N and NO3--N) and available P compared to those in natural marshes and tidal flats. The distribution of soil TC, TN, and TP demonstrated greater spatial heterogeneity in natural marshes and tidal flats, while the converted areas were more uniform and became hotspots for N and P accumulation. Coastal conversion altered soil C:N:P stoichiometry, with cropland soils exhibiting a lower N:P ratio (2.9 ± 1.1), indicating that long-term application of N and P fertilizers could decrease the N:P ratio, as P is more retained in the soil than N. Furthermore, it was observed that the dynamics of C, N and P, as well as their stoichiometry, are closely linked to soil physicochemical properties, especially soil organic matter and texture. These findings highlight that coastal conversion and associated management practices markedly affected soil C, N and P dynamics in a representative wetland area of the subtropical regions, leading to a reshaping of their stoichiometric balances, particularly in the topsoil layer.

Linking soil phosphorus availability and phosphatase functional genes to coastal marsh erosion: Implications for nutrient cycling and wetland restoration.

Accelerated marsh erosion caused by climate change and human activity may have important implications for nutrient cycling and availability. However, how erosion affects phosphorus (P) transformation and microbial function in subtropical coastal marshes remains largely unknown. Here we assessed soil P fractions, availability and the phoD-harboring bacterial community along a marsh erosion gradient (non-eroded, lightly eroded, and heavily eroded). We showed that marsh erosion caused a shift in P fractions, leading to a decrease in P availability and a reduction in concentrations of labile P, moderately labile P, and stable P by 20 %, 9 %, and 17 % respectively. The abundance and diversity of phoD phosphatase genes decreased dramatically along the erosion gradient and were lower at heavily eroded sites than at non-eroded sites. Marsh erosion reshaped phoD gene community composition, and Corallococcus, Amycolatopsis, and Phaeobacter were identified as the dominant phoD-harboring microbes. Notably, marsh erosion reduced the complexity and stability of the phoD-harboring bacterial network, and heavily eroded sites have fewer network edges and nodes than non-eroded sites. The dynamics of soil P fractions, availability, and phoD-harboring bacterial communities driven by marsh erosion are largely shaped by substrate availability and soil properties (e.g., nutrients, pH, electrical conductivity, and moisture). Additionally, strong linkages between P availability and the abundance and diversity of phosphatase genes following erosion, suggest that phosphatase drives P mineralization and dissolution, and erosion weakens the regulation of P transformation by reshaping the phoD phosphatase gene community. Our findings indicate that marsh erosion alters soil P fractions and phoD-harboring bacterial communities, which weakens microbial regulation of P transformation and availability, thereby significantly reducing soil P pools and availability. Our findings broaden understanding of the impacts of coastal erosion on nutrient balance and ecosystem function, offering valuable perspectives that could inform wetland restoration and coastal management strategies.

Coastal wetland conversion to aquaculture pond reduced soil P availability by altering P fractions, phosphatase activity, and associated microbial properties.

Reclamation and conversion of wetlands strongly affect nutrient cycling and ecosystem functions, while little attention has been paid to the effects of converting coastal wetland to aquaculture on the cycling and balance of soil phosphorus (P). Herein, we investigated soil P fractions, alkaline phosphatase (ALP) activity, and associated microbial properties following coastal wetland conversion in subtropical China. Soil P availability (especially resin-P) concentration and ALP activity in wetland were significantly higher than those in pond. The conversion of coastal wetlands to aquaculture significantly reduced the abundance and diversity of bacterial phoD genes and altered their community structure. The lower phosphatase activity and associated microbial properties after wetland conversion suggested a weaker capacity of microbes to transform organic P (Po) to inorganic P (Pi), consistent with the low P availability but the high Po:Pi ratio in pond. Structural equation modeling indicated that the conversion of the wetland to the pond decreased ALP activity and P availability by affecting soil variables such as bulk density, pH, the carbon: nitrogen ratio, and/or moisture. It was concluded that wetland conversion to pond reduced soil P availability and phosphatase activity, altered the abundance, diversity and community composition of the phoD gene, and ultimately affected coastal P cycles and balances. Moreover, an extended corollary is that the smaller amounts of variation in soil total P and lower labile P concentrations in pond than in wetland suggest that large amounts of P (introduced in feed and not harvested in shrimp) are "lost" from the system. Thus, aquaculture ponds might serve as a source of P for the surrounding environment. More investigations focusing on the P biogeochemical cycle and its potential impacts on adjacent ocean environments at regional and global scales is urgently needed, which could contribute to better management of coastal land uses.

Substantial increase in P release following conversion of coastal wetlands to aquaculture ponds from altered kinetic exchange and resupply capacity.

The reclamation of wetlands and its subsequent conversion to aquaculture may alter regional nutrient (im)mobilization and cycling, although direct assessments of phosphorus (P) cycling and its budget balance following wetland conversion are currently scarce. Here, parallel field experiments were conducted to investigate and compare the availability and mobilization mechanisms of P from natural coastal wetlands and the adjacent converted aquaculture ponds based on high-resolution diffusive gradient in thin films (DGT) and dialysis (HR-Peeper) techniques and the DGT-induced fluxes in sediments (DIFS) model. The study found that the conversion of wetland to pond strongly reduced the sediment P pool by changing its forms and distribution. High-resolution data showed that concentrations of labile P and soluble reactive P across the sediment-water profiles were markedly enhanced by the converted aquaculture pond, although they exhibited large spatiotemporal heterogeneity. Moreover, the synchronous distribution of labile P, iron (Fe) and sulfur (S) across profiles in coastal wetlands indicated that the dissolution of Fe (III) oxyhydroxide-phosphate complexes coupled with sulfate reduction were the main mechanisms regulating sediment P mobilization in coastal areas. However, the converted aquaculture pond weakened or even reversed this dependence by decoupling the Fe-S-P reactions by changing the sediment structure and nutrient balance. Substantial increases in labile P, Fe and S fluxes in the pond suggested the conversion of wetland to aquaculture facilitated the internal release of P, Fe and S from sediment into water. The high resupply parameter (R) and desorption rate (k-1) combined with low response time (Tc) in the pond, as fitted by DIFS model, indicated the strong resupply capacity and fast kinetic exchange of sediment P across the sediment-water interface, which is consistent with the high P diffusion fluxes recorded in the pond. It was concluded that converted aquaculture ponds act as an important source of P release in coastal areas, potentially exacerbating water quality degradation and eutrophication. Specific initiatives and actions are therefore urgently needed to alleviate the internal P-loading during aquaculture.

Biogeochemical behavior of P in the soil and porewater of a low-salinity estuarine wetland: Availability, diffusion kinetics, and mobilization mechanism.

Estuarine wetlands, which typically store large amounts of phosphorus (P), are experiencing increased salinity as well as changed environmental factors caused by rising sea levels. In this study, the seasonal dynamics of P speciation, availability, and biogeochemical couplings with iron (Fe)-sulfur (S) in soil and porewater were measured in a low-salinity estuarine wetland using in situ high-resolution diffusive gradients in thin films (DGT) and dialysis (HR-Peeper) techniques. The diffusion kinetics and resupply capacity of P from the soil phase to solution were simulated using a DGT-induced fluxes in soils (DIFS) model. The transition from freshwater to brackish wetlands reduced soil P pools and shifted to more recalcitrant speciation. The concentration of DGT-labile P across the soil-water profiles ranged from 0.002 to 0.039 (mean: 0.015) mg L-1, which increased with increasing salinity in both the field and mesocosm experiments. The distributions of labile and soluble P showed high heterogeneity across the profiles, and there were some sharp peak values below the soil-water interface (SWI), which significantly increased the concentration and lability of P. The strong coupling between labile P and Fe (S) provided direct evidence for the coexistence of iron reduction (IR) and sulfate reduction (SR) in the estuary, while IR might predominate in P mobilization in the brackish environment because of higher labile Fe concentrations and stronger Fe-P couplings. The diffusion fluxes of P were positive at both sites, demonstrating that the kinetics of P were from the soils to the overlying water. Higher R and k-1 values fitted in the DIFS model implied that a stronger resupply capacity and desorption rate and thus faster remobilization kinetics of P occurred with increasing salinity. Our findings indicated that increased salinity (even at low levels) can alter the desorption rate and resupply capacity of soil P in estuarine wetlands and accelerate P remobilization and release by regulating the IR and SR processes, thereby leading to the deterioration of water quality.

Effects of wetland types on dynamics and couplings of labile phosphorus, iron and sulfur in coastal wetlands during growing season.

Wetland type plays an important role in controlling the phosphorus (P) biogeochemical cycle, while its effect on labile P dynamics and coupling with iron (Fe) and sulfur (S) in coastal wetlands remains unclear. In this study, chemical sequential extraction and high-resolution diffusive gradients in thin-film (DGT) techniques were employed to investigate P forms, mobilization, and labile Fe-S-P coupling in several coastal wetland types [i.e., natural wetland (NW), aquaculture pond (AP), artificial (ARW) and natural restored wetlands (NRW)]. Compared with NW, AP decreased the total P by 40.6%. The concentrations of soil organic P and inorganic P (including NaOH-extractable P and HCl-extractable P) were significantly increased in ARW, but decreased in AP and NRW. DGT-labile P, Fe, and S concentrations changed significantly in different wetland types, and the labile P concentrations in AP were significantly higher than those in the others. Similar spatial distribution dynamics and significant positive relationships between labile P, Fe, and S concentrations in NW and AP confirmed that intense reduction in iron and sulfate are the key mechanisms regulating P mobilization. However, these relationships were decoupled in restored wetlands, suggesting that the Fe redox-coupled P mobilization and sulfate reduction were sensitive to wetland changes. The diffusion fluxes of P across the soil-water interface were positive in AP (0.619 pg·cm-2·s-1), indicating that P was released from soil to the overlying-water. We concluded that coastal wetland types altered soil P forms, availability, and labile Fe-S-P coupling, and the natural restored wetland could help stabilize the soil P pool and eventually controlled the mobilization and release of P.