Identification and analysis of short-term and long-term salt-associated lncRNAs in the leaf of Avicennia marina.

As a highly salt-resistant mangrove, Avicennia marina can thrive in the hypersaline water. The leaves of Avicennia marina play a crucial role in salinity stress adaptability by secreting salt. Although the functions of long non-coding RNAs (lncRNAs) in leaves remain unknown, they have emerged as regulators in leaf development, aging and salt response. In this study, we employed transcriptomic data of both short-term and long-term salt treated leaves to identify salt-associated lncRNAs of leaf tissue. As a result, 687 short-term and 797 long-term salt-associated lncRNAs were identified. Notably, both short-term and long-term salt-associated lncRNAs exhibited slightly longer lengths and larger exons, but smaller introns compared with salt-non-associated lncRNAs. Furthermore, salt-associated lncRNAs also displayed higher tissue-specificity than salt-non-associated lncRNAs. Most of the salt-associated lncRNAs were common to short- and long-term salt treatments. And about one fifth of the downregulated salt-associated lncRNAs identified both in two terms were leaf tissue-specific lncRNAs. Besides, these leaf-specific lncRNAs were found to be involved in the oxidation-reduction and photosynthesis processes, as well as several metabolic processes, suggesting the noticeable functions of salt-associated lncRNAs in regulating salt responses of Avicennia marina leaves.

High temperature and eutrophication alter biomass allocation of black mangrove (Avicennia germinans L.) seedlings.

Mangrove restoration is underway along tropical coastlines to combat their rapid worldwide decline. However, restoration success is limited due to local drivers such as eutrophication, and global drivers such as climate change, yet their interactions remain unclear. We conducted a mesocosm experiment to assess the impact of increased nutrients and temperature on the photosynthetic efficiency and development of black mangrove seedlings. Seedlings exposed to high temperature and eutrophication showed reduced root growth and disproportionally long stems, with lower net assimilation rates. This architectonical imbalance between root and stem growth may increase susceptibility to physical disturbances and dislodgement. Notably, none of the experimental seedlings displayed signs of photophysiological stress, and those exposed to increased nutrients and temperature exhibited robust photosynthetic performance. The disbalance in biomass allocation highlights the importance of considering local nutrient status and hydrodynamic conditions in restoration projects, ensuring the effective anchorage of mangrove seedlings and restoration success under a warming climate.

Widespread retreat of coastal habitat is likely at warming levels above 1.5 °C.

Several coastal ecosystems-most notably mangroves and tidal marshes-exhibit biogenic feedbacks that are facilitating adjustment to relative sea-level rise (RSLR), including the sequestration of carbon and the trapping of mineral sediment1. The stability of reef-top habitats under RSLR is similarly linked to reef-derived sediment accumulation and the vertical accretion of protective coral reefs2. The persistence of these ecosystems under high rates of RSLR is contested3. Here we show that the probability of vertical adjustment to RSLR inferred from palaeo-stratigraphic observations aligns with contemporary in situ survey measurements. A deficit between tidal marsh and mangrove adjustment and RSLR is likely at 4 mm yr-1 and highly likely at 7 mm yr-1 of RSLR. As rates of RSLR exceed 7 mm yr-1, the probability that reef islands destabilize through increased shoreline erosion and wave over-topping increases. Increased global warming from 1.5 °C to 2.0 °C would double the area of mapped tidal marsh exposed to 4 mm yr-1 of RSLR by between 2080 and 2100. With 3 °C of warming, nearly all the world's mangrove forests and coral reef islands and almost 40% of mapped tidal marshes are estimated to be exposed to RSLR of at least 7 mm yr-1. Meeting the Paris agreement targets would minimize disruption to coastal ecosystems.

Foliar water uptake enables embolism removal in excised twigs of Avicennia marina.

Embolism refilling is thought to require relaxation of xylem tension, and it is unclear whether and how tall trees or plants growing in arid or saline soils recover from embolism. We tested whether foliar water uptake could enable embolism refilling in dehydrated twigs of the grey mangrove (Avicennia marina). Four dehydrated twigs were imaged by laboratory-based micro-computed tomography before and after wetting leaves. Emboli were observed in dehydrated stems and leaves. Embolism decreased with increasing distance from the cut end of stems, suggesting that stem emboli were caused by cutting. A significant (P = 0.026) c. 80% reduction in the embolised area was observed in leaves between the start and the end of the experiment (29 ± 10 h after wetting). Embolus diameter was unaffected by wetting. Embolism refilling occurred slowly, in stems embolised by cutting and leaves embolised by cutting and/or dehydration. The lack of response of embolus diameter to wetting suggests that capillarity was not the main mechanism for refilling. Results show that excised twigs of A. marina are able to recover from embolism by absorption of atmospheric water and call for studies under natural conditions.

Regulation of a Cytochrome P450 Gene CYP94B1 by WRKY33 Transcription Factor Controls Apoplastic Barrier Formation in Roots to Confer Salt Tolerance.

Salinity is an environmental stress that causes decline in crop yield. Avicennia officinalis and other mangroves have adaptations such as ultrafiltration at the roots aided by apoplastic cell wall barriers to thrive in saline conditions. We studied a cytochrome P450 gene from A. officinalis, AoCYP94B1, and its putative ortholog in Arabidopsis (Arabidopsis thaliana), AtCYP94B1, which are involved in apoplastic barrier formation. Both genes were induced by 30 min of salt treatment in the roots. Heterologous expression of AoCYP94B1 in the atcyp94b1 Arabidopsis mutant and wild-type rice (Oryza sativa) conferred increased NaCl tolerance to seedlings by enhancing root suberin deposition. Histochemical staining and gas chromatography-tandem mass spectrometry quantification of suberin precursors confirmed the role of CYP94B1 in suberin biosynthesis. Using chromatin immunoprecipitation and yeast one-hybrid and luciferase assays, we identified AtWRKY33 as the upstream regulator of AtCYP94B1 in Arabidopsis. In addition, atwrky33 mutants exhibited reduced suberin and salt-sensitive phenotypes, which were rescued by expressing 35S::AtCYP94B1 in the atwrky33 background. This further confirmed that AtWRKY33-mediated regulation of AtCYP94B1 is part of the salt tolerance mechanism. Our findings may help efforts aimed at generating salt-tolerant crops.

Molecular cloning and expression of AmCDPK from mangrove Avicennia marina under elevated temperature.

Considered as an essential calcium sensor, the calcium-dependent protein kinase (CDPK) family plays a critical part in terrestrial plants' responses to both biotic and abiotic stresses. In the study, Avicennia marina was proved to have better heat tolerance than other species. A CDPK gene was cloned from mangrove species A. marina using RACE-PCR and designated as AmCDPK. By predicting and analyzing its properties, structures and expression patterns, we found that the amino acid sequence, containing a kinase domain and four EF-hand Ca2+-binding sites, shared high identity with Handroanthus impetiginosus and Sesamum indicum. Quantitative real-time PCR data analysis suggested that AmCDPK demonstrated significant up-regulation under heat stress. It is likely that AmCDPK is a versatile gene involved in various stresses, including dehydration, cold, light, defense and ABA stress responses by analyzing cis-elements. It is the first time that CDPKs from mangroves have been cloned and our results brought evidence to the effect of AmCDPK on heat stress, which is particularly important under the background of global warming.

Proline metabolism and molecular cloning of AmP5CS in the mangrove Avicennia marina under heat stress.

Proline is one of the most important compatible osmolyte in cells, which accumulates in response to various stresses, including salt, water deficit, heavy metal, pathogen infection and extreme temperature. In this study, a growth chamber was employed to simulate heat environment for Avicennia marina seedlings. We detected some physiological indices in the leaves of A. marina at 40 °C, including the activity of delta-1-pyrroline-5-carboxylate synthase (P5CS), the content of free proline and soluble protein, transpiration rate and membrane permeability, and discussed the relationship between these five indices and heat resistant ability. And then a P5CS gene was cloned from A. marina using homologous cloning and rapid amplification of cDNA ends methods. It was designated as AmP5CS, encoding a protein that contained a feedback inhibition site of proline, proA, proB, conserved Leu zipper, GSA-DH domain and other functional domains of P5CS protein in high plants. Expression analysis of AmP5CS gene indicated it was involved in heat stress response. It is the first time that P5CS from A. marina has been cloned and the findings laid the foundation of figuring out heat resistant mechanisms and relieving heat damage, which is significant during global warming.

Salt tolerance and exclusion in the mangrove plant Avicennia marina in relation to root apoplastic barriers.

Salt tolerance and the possible functions of suberization on salt exclusion and secretion were examined in a dominant mangrove plant, Avicennia marina. The results showed that low salinities (10‰ and 20‰) almost has no negative effect on A. marina, however significant growth inhibitions were observed in the seedlings grown in higher salinities (30‰ and 40‰). With the increases of salinity, increased tissue Na+ content and enhanced salt secretion by glands were observed. Obvious suberization thickening were detected both in the exodermis and endodermis of the roots after salt pretreatment when compared to the roots without salt treatment. More importantly, the present data further confirmed that these root apoplastic barriers would directly decrease Na+ loading into xylem. Higher salt tolerance was observed in the seedlings pre-cultivated by salty tide when compared to fresh water cultivated A. marina. In summary, this study suggests a barrier property of suberization in dealing with salt exclusion in mangroves, a moderate salt pre-treatment may benefit plant withstanding high salinity.

Night and day: Shrinking and swelling of stems of diverse mangrove species growing along environmental gradients.

Tree stems swell and shrink daily, which is thought to reflect changes in the volume of water within stem tissues. We observed these daily patterns using automatic dendrometer bands in a diverse group of mangrove species over five mangrove forests across Australia and New Caledonia. We found that mangrove stems swelled during the day and shrank at night. Maximum swelling was highly correlated with daily maxima in air temperature. Variation in soil salinity and levels of tidal inundation did not influence the timing of stem swelling over all species. Medium-term increases in stem circumference were highly sensitive to rainfall. We defoliated trees to assess the role of foliar transpiration in stem swelling and shrinking. Defoliated trees showed maintenance of the pattern of daytime swelling, indicating that processes other than canopy transpiration influence the temporary stem diameter increments, which could include thermal swelling of stems. More research is required to understand the processes contributing to stem shrinking and swelling. Automatic Dendrometer Bands could provide a useful tool for monitoring the response of mangroves to extreme climatic events as they provide high-frequency, long-term, and large-scale information on tree water status.

Shoot surface water uptake enables leaf hydraulic recovery in Avicennia marina.

The significance of shoot surface water uptake (SSWU) has been debated, and it would depend on the range of conditions under which it occurs. We hypothesized that the decline of leaf hydraulic conductance (Kleaf ) in response to dehydration may be recovered through SSWU, and that the hydraulic conductance to SSWU (Ksurf ) declines with dehydration. We quantified effects of leaf dehydration on Ksurf and effects of SSWU on recovery of Kleaf in dehydrated leaves of Avicennia marina. SSWU led to overnight recovery of Kleaf , with recovery retracing the same path as loss of Kleaf in response to dehydration. SSWU declined with dehydration. By contrast, Ksurf declined with rehydration time but not with dehydration. Our results showed a role of SSWU in the recovery of leaf hydraulic conductance and revealed that SSWU is sensitive to leaf hydration status. The prevalence of SSWU in vegetation suggests an important role for atmospheric water sources in maintenance of leaf hydraulic function, with implications for plant responses to changing environments.

Interactive effects of temperature and nutrients on mangrove seedling growth and implications for establishment.

The establishment and wellbeing of seedlings governs the spread and survival of mangrove forests. Eutrophication and global warming are major challenges endangering mangrove ecosystem integrity. How these stressors affect seedling growth is not well understood. In a mesocosm experiment we grew mangrove seedlings in temperature-controlled chambers and investigated single and combined effects of temperature (23 and 33 °C), organic matter and dissolved nutrients on seedling trait morphology. Seedling survival was lowest in organic matter treatments. Combined effects of temperature and nutrients caused significant differences in root morphology with fewer but longer and thicker 3rd order roots, fewer 2nd and no 1st order roots in nutrient-enriched (23 °C) compared to non-enriched treatments (33 °C). Our results indicate these seedlings are less resilient to withstand their dynamic environment, in which they must settle and establish, due to lower root complexity. Mangrove ecosystems are negatively affected by global and local stresses; if new seedlings, which support forest recovery, are also affected then this amplifies stresses.

A general framework for propagule dispersal in mangroves.

Dispersal allows species to shift their distributions in response to changing climate conditions. As a result, dispersal is considered a key process contributing to a species' long-term persistence. For many passive dispersers, fluid dynamics of wind and water fuel these movements and different species have developed remarkable adaptations for utilizing this energy to reach and colonize suitable habitats. The seafaring propagules (fruits and seeds) of mangroves represent an excellent example of such passive dispersal. Mangroves are halophytic woody plants that grow in the intertidal zones along tropical and subtropical shorelines and produce hydrochorous propagules with high dispersal potential. This results in exceptionally large coastal ranges across vast expanses of ocean and allows species to shift geographically and track the conditions to which they are adapted. This is particularly relevant given the challenges presented by rapid sea-level rise, higher frequency and intensity of storms, and changes in regional precipitation and temperature regimes. However, despite its importance, the underlying drivers of mangrove dispersal have typically been studied in isolation, and a conceptual synthesis of mangrove oceanic dispersal across spatial scales is lacking. Here, we review current knowledge on mangrove propagule dispersal across the various stages of the dispersal process. Using a general framework, we outline the mechanisms and ecological processes that are known to modulate the spatial patterns of mangrove dispersal. We show that important dispersal factors remain understudied and that adequate empirical data on the determinants of dispersal are missing for most mangrove species. This review particularly aims to provide a baseline for developing future research agendas and field campaigns, filling current knowledge gaps and increasing our understanding of the processes that shape global mangrove distributions.

Growth and antioxidative response of two mangrove plants to interaction between aquaculture effluent and BDE-99.

Mangroves are subject to contamination of polybrominated diphenyl ethers (PBDEs) due to waste and wastewater disposal, and aquaculture effluent (AE) from nearby aquaculture activities. However, the response of mangrove plants to these two stresses and their interaction has seldom been reported. A six-month microcosm study, planted with either Kandelia obovata (Ko) or Avicennia marina (Am), the two most dominant species in South China mangrove swamps, was conducted to investigate the effects of BDE-99, and the interactions of BDE-99 (one of the most abundant PBDE congeners) and AE on growth and physiological responses of these plants. In addition to mixed stressors, both stressors were also applied individually. Results showed that Avicennia was more tolerant to BDE-99 contamination than Kandelia, as reflected by the reduced biomass, but increased superoxide radical (O2-⁎) release and malondialdehyde (MDA) content in Kandelia. Addition of AE alleviated toxicity of BDE-99 in Kandelia by promoting biomass but lowering oxidative stress and MDA production. The hormesis model also demonstrated that the interaction between BDE-99 and AE on leaf and root MDA and O2-⁎ content in both Kandelia and Avicennia were mostly antagonistic. Activities of catalase (CAT), superoxide dismutase (SOD) and peroxidase (POD) in both leaf and root of Kandelia were reduced by BDE-99. On the contrary, BDE-99 significantly enhanced the three enzyme activities in Avicennia root at month 3. Addition of AE also significantly enhanced root CAT, POD and SOD activities, and leaf SOD in both plant species to remove excess ROS produced under BDE-99 exposure. These results indicated that the tolerance of mangrove plants to oxidative stresses depended on antioxidative enzymes that were inducible.

Modeling multi-decadal mangrove leaf area index in response to drought along the semi-arid southern coasts of Iran.

Leaf Area Index (LAI; as an indicator of the health) of the mangrove ecosystems on the northern coasts of the Persian Gulf and the Gulf of Oman was measured in the field and modeled in response to observed (1986-2017) and predicted (2018-2100) drought occurrences (quantified using the Standardized Precipitation Index [SPI]). The relationship of LAI with the normalized difference vegetation index (NDVI) obtained from satellite images was quantified, the LAI between 1986 and 2017 retrospectively estimated, and a relationship between LAI and SPI developed for the same period. Long-term climate data were used as input in the RCP8.5 climate change scenario to reconstruct recent and forecast future drought intensities. Both the NDVI and the SPI were strongly related with the LAI, indicating that realistic LAI values were derived from historic satellite data to portray annual changes of LAI in response to changes in SPI. Our findings show that projected future drought intensities modeled by the RCP8.5 scenario increase more and future LAIs decreased more on the coasts of the Gulf of Oman than the coasts of the Persian Gulf in the coming decades. The year 1998 was the most significant change-point for mean annual rainfall amounts and drought occurrences as well as for LAIs and at no time between 1998 and 2017 or between 2018 and 2100 are SPI and LAI values expected to return to pre-1998 values. LAI and SPI are projected to decline sharply around 2030, reach their lowest levels between 2040 and 2070, and increase and stabilize during the late decades of the 21st century at values similar to the present time. Overall, this study provides a comprehensive picture of the responses of mangroves to fluctuating future drought conditions, facilitating the development of management plans for these vulnerable habitats in the face of future climate change.

Tropicalization of the barrier islands of the northern Gulf of Mexico: A comparison of herbivory and decomposition rates between smooth cordgrass (Spartina alterniflora) and black mangrove (Avicennia germinans).

The expansion of black mangrove Avicennia germinans into historically smooth cordgrass Spartina alterniflora-dominated marshes with warming temperatures heralds the migration of the marsh-mangrove ecotone northward in the northern Gulf of Mexico. With this shift, A. germinans is expected to outcompete S. alterniflora where it is able to establish, offering another prevalent food source to first order consumers. In this study, we find A. germinans leaves to be preferable to chewing herbivores, but simultaneously, chewing herbivores cause more damage to S. alterniflora leaves. Despite higher nitrogen content, A. germinans leaves decomposed slower than S. alterniflora leaves, perhaps due to other leaf constituents or a different microbial community. Other studies have found the opposite in decomposition rates of the two species' leaf tissue. This study provides insights into basic trophic process, herbivory and decomposition, at the initial stages of black mangrove colonization into S. alterniflora salt marsh.

Global-scale dispersal and connectivity in mangroves.

Dispersal provides a key mechanism for geographical range shifts in response to changing environmental conditions. For mangroves, which are highly susceptible to climate change, the spatial scale of dispersal remains largely unknown. Here we use a high-resolution, eddy- and tide-resolving numerical ocean model to simulate mangrove propagule dispersal across the global ocean and generate connectivity matrices between mangrove habitats using a range of floating periods. We find high rates of along-coast transport and transoceanic dispersal across the Atlantic, Pacific, and Indian Oceans. No connectivity is observed between populations on either side of the American and African continents. Archipelagos, such as the Galapagos and those found in Polynesia, Micronesia, and Melanesia, act as critical stepping-stones for dispersal across the Pacific Ocean. Direct and reciprocal dispersal routes across the Indian Ocean via the South Equatorial Current and seasonally reversing monsoon currents, respectively, allow connectivity between western Indian Ocean and Indo-West Pacific sites. We demonstrate the isolation of the Hawaii Islands and help explain the presence of mangroves on the latitudinal outlier Bermuda. Finally, we find that dispersal distance and connectivity are highly sensitive to the minimum and maximum floating periods. We anticipate that our findings will guide future research agendas to quantify biophysical factors that determine mangrove dispersal and connectivity, including the influence of ocean surface water properties on metabolic processes and buoyancy behavior, which may determine the potential of viably reaching a suitable habitat. Ultimately, this will lead to a better understanding of global mangrove species distributions and their response to changing climate conditions.

The response of stocks of C, N, and P to plant invasion in the coastal wetlands of China.

The increasing success of invasive plant species in wetland areas can threaten their capacity to store carbon, nitrogen, and phosphorus (C, N, and P). Here, we have investigated the relationships between the different stocks of soil organic carbon (SOC), and total C, N, and P pools in the plant-soil system from eight different wetland areas across the South-East coast of China, where the invasive tallgrass Spartina alterniflora has replaced the native tall grasses Phragmites australis and the mangrove communities, originally dominated by the native species Kandelia obovata and Avicennia marina. The invasive success of Spartina alterniflora replacing Phragmites australis did not greatly influence soil traits, biomass accumulation or plant-soil C and N storing capacity. However, the resulting higher ability to store P in both soil and standing plant biomass (approximately more than 70 and 15 kg P by ha, respectively) in the invasive than in the native tall grass communities suggesting the possibility of a decrease in the ecosystem N:P ratio with future consequences to below- and aboveground trophic chains. The results also showed that a future advance in the native mangrove replacement by Spartina alterniflora could constitute a serious environmental problem. This includes enrichment of sand in the soil, with the consequent loss of nutrient retention capacity, as well as a sharp decrease in the stocks of C (2.6 and 2.2 t C ha-1 in soil and stand biomass, respectively), N, and P in the plant-soil system. This should be associated with a worsening of the water quality by aggravating potential eutrophication processes. Moreover, the loss of carbon and nutrient decreases the potential overall fertility of the system, strongly hampering the reestablishment of woody mangrove communities in the future.

A facultative mutualistic feedback enhances the stability of tropical intertidal seagrass beds.

Marine foundation species such as corals, seagrasses, salt marsh plants, and mangrove trees are increasingly found to engage in mutualistic interactions. Because mutualisms by their very nature generate a positive feedback between the species, subtle environmental impacts on one of the species involved may trigger mutualism breakdown, potentially leading to ecosystem regime shifts. Using an empirically parameterized model, we investigate a facultative mutualism between seagrass and lucinid bivalves with endosymbiotic sulfide-oxidizing gill bacteria in a tropical intertidal ecosystem. Model predictions for our system show that, by alleviating the build-up of toxic sulfide, this mutualism maintains an otherwise intrinsically unstable seagrass ecosystem. However, an increase in seagrass mortality above natural levels, due to e.g. desiccation stress, triggers mutualism breakdown. This pushes the system in collapse-and-recovery dynamics ('slow-fast cycles') characterized by long-term persistent states of bare and seagrass-dominated, with rapid transitions in between. Model results were consistent with remote sensing analyses that suggest feedback-mediated state shifts induced by desiccation. Overall, our combined theoretical and empirical results illustrate the potential of mutualistic feedbacks to stabilize ecosystems, but also reveal an important drawback as small environmental changes may trigger shifts. We therefore suggest that mutualisms should be considered for marine conservation and restoration of seagrass beds.

Effects of elevated atmospheric CO2 and increased tidal flooding on leaf gas-exchange parameters of two common mangrove species: Avicennia marina and Rhizophora stylosa.

In this study, we examined interactive effects of elevated atmospheric CO2, concentrations, and increased tidal flooding on two mangroves species, Avicennia marina and Rhizophora stylosa. Leaf gas-exchange parameters (photosynthesis, transpiration rates, water-use efficiency, stomatal conductance, and dark respiration rates) were measured monthly on more than 1000 two-year-old seedlings grown in greenhouses for 1 year. In addition, stomatal density and light curve responses were determined at the end of the experiment. Under elevated CO2 concentrations (800 ppm), the net photosynthetic rates were enhanced by more than 37% for A. marina and 45% for R. stylosa. This effect was more pronounced during the warm season, suggesting that an increase in global temperatures would further enhance the photosynthetic response of the considered species. Transpiration rates decreased by more than 15 and 8% for A. marina and R. stylosa, respectively. Consequently, water-use efficiency increased by 76% and 98% for A. marina and R. stylosa, respectively, for both species, which will improve drought resistance. These responses to elevated CO2 were minimized (by 5%) with longer flooding duration. Consequently, future increases of atmospheric CO2 may have a strong and positive effect on juveniles of A. marina and R. stylosa during the next century, which may not be suppressed by the augmentation of tidal flooding duration induced by sea-level rise. It is possible that this effect will enhance seedling dynamic by increasing photosynthesis, and therefore will facilitate their settlements in new area, extending the role of mangrove ecosystems in carbon sequestration and climate change mitigation.

Proteomic analysis on mangrove plant Avicennia marina leaves reveals nitric oxide enhances the salt tolerance by up-regulating photosynthetic and energy metabolic protein expression.

Avicennia marina (Forsk.) Vierh is one of the most salt-tolerant mangrove species. Our previous study revealed that nitric oxide (NO) enhanced the salt tolerance of A. marina by promoting salt secretion and Na+ sequestration under salt stress. However, little is known about the regulation of NO on proteomic profiling for this mangrove species. In this study, we used sodium nitroprusside (SNP), an NO donor, to investigate the regulatory mechanism of NO on salt tolerance of A. marina according to physiological and proteomic aspects. Photosynthesis data showed that the reduction in photosynthesis caused by high salinity treatment (400 mM NaCl) could be partially recovered by addition of SNP (100 µM). Further analysis revealed that the high salinity treatment could induce not only the stomatal limitation but also non-stomatal limitation on photosynthetic reduction, while SNP addition could restore the non-stomatal limitation, implying that the application of SNP was beneficial to the metabolic process in leaves. Proteomic analysis identified 49 differentially expressed proteins involved in various biological processes such as photosynthesis, energy metabolism, primary metabolism, RNA transcription, protein translation and stress response proteins. Under high salinity treatment, the abundances of proteins related to photosynthesis, such as ribulose-phosphate 3-epimerase (RPE, spot 3), RuBisCO large subunit (RBCL, spot 4, 5, 24), RuBisCO activase A (RCA, spot 17, 18) and quinine oxidoreductase-like protein isoform 1 (QOR1, spot 23), were significantly decreased. However, the abundance of proteins such as RBCL (spot 5, 9) and QOR1 (spot 23) were increased by SNP addition. In addition, exogenous NO supply alleviated salt tolerance by increasing the accumulation of some proteins involved in energy metabolism (spot 15), primary metabolism (spot 25, 45, 46), RNA transcription (spot 36) and stress response proteins (spot 12, 21, 26, 37, 43). The transcriptional levels of nine selected proteins were mostly consistent with their protein abundance except spot 46. Overall, the presented data demonstrated that NO has a positive effect on improving salt tolerance in A. marina by regulating the protein abundance involved in photosynthesis, energy metabolism, primary metabolism and stress response.