Diagnostic species are crucial for the functioning of plant associations in inland salt marshes.

Salt marsh vegetation is considered unique and valuable and has been legally protected in Europe for years but is still declining. Its protection is related to vegetation syntaxonomical units. The characteristic combination of diagnostic species is used to create this syntaxonomical system. The aim of our novel study was to assess whether diagnostic species are sufficient for characterising vegetation functioning. Moreover, we included biochemical traits not considered to date in vegetation ecology. We hypothesised that (1) diagnostic species are crucial for the functioning of inland salt marsh vegetation and (2) their morphological and biochemical traits define the functioning of typical salt marsh associations. We chose three typical inland associations to test our hypotheses and measured the morphological and biochemical functional traits of their diagnostic plant species. Our research has shown that diagnostic species play a crucial role not only in distinguishing typical inland salt marsh associations but also in determining their functioning. Among the analysed associations, Salicornietum ramosissimae was the most adaptable to osmotic and oxidative stress under soil salinity. Triglochino maritimae-Glaucetum maritimae showed the lowest salt resistance, as indicated by the highest osmotic and oxidative stress and stress responses. Our findings may facilitate the practical application of new approaches and protection strategies for inland salt marsh habitats.

ROS Homeostasis and Antioxidants in the Halophytic Plants and Seeds.

Reactive oxygen species (ROS) are excited or partially reduced forms of atmospheric oxygen, which are continuously produced during aerobic metabolism like many physiochemical processes operating throughout seed life. Previously, it was believed that ROS are merely cytotoxic molecules, however, now it has been established that they perform numerous beneficial functions in plants including many critical roles in seed physiology. ROS facilitate seed germination via cell wall loosening, endosperm weakening, signaling, and decreasing abscisic acid (ABA) levels. Most of the existing knowledge about ROS homeostasis and functions is based on the seeds of common plants or model ones. There is little information about the role of ROS in the germination process of halophyte seeds. There are several definitions for halophytic plants, however, we believed "halophytes are plants that can grow in very saline environment and complete their life cycle by adopting various phenological, morphological and physiological mechanisms at canopy, plant, organelle and molecular scales". Furthermore, mechanisms underlying ROS functions such as downstream targets, cross-talk with other molecules, and alternative routes are still obscure. The primary objective of this review is to decipher the mechanisms of ROS homeostasis in halophytes and dry seeds, as well as ROS flux in germinating seeds of halophytes.

Technogenic soil salinisation, vegetation, and management shape microbial abundance, diversity, and activity.

The importance of the microbiome in the functioning of degraded lands in industrialised zones is significant. However, little is known about how environmental parameters affect microbial abundance, structure, diversity, and especially specific guilds involved in the nitrogen cycle in saline soils influenced by the soda industry. To address this knowledge gap, our research focused on assessing the microbiota in relation to soil properties and plant species composition across two transects representing different types of land use: saline wasteland and arable fields. Our findings show that the microbial communities were the most affected not only by soil salinity but also by pH and the composition of plant species. Taxonomic variability was the most shaped by salinity together with management type and CaCO3 content. The impact of salinity on the soil microbiome was manifested in a reduced abundance of bacteria and fungi, a lower number of observed phylotypes, reduced modularity, and a lower abundance of the nitrifying guild. Denitrification and nitrogen fixation were less affected by salinity. The last process was correlated with calcium carbonate. CaCO3 was also associated with microbial taxonomic variability and the overall microbial activity caused by hydrolases, which could aid organic matter turnover in saline but carbonate-rich sites. Bacterial genera such as Bacillus, Peanibacillus, and Rhodomicrobium, in addition to fungal taxa such as Cadophora, Mortierella globalpina, Preussia flanaganii, and Chrysosporium pseudomerdarium, show potential as favourable candidates for possible bioremediation initiatives. These results can be applied to future land reclamation projects. FUNDING INFORMATION: This research received no specific grant from funding agencies in the public, commercial, or not-for-profit sectors.

The utility of airborne hyperspectral and satellite multispectral images in identifying Natura 2000 non-forest habitats for conservation purposes.

Aerial hyperspectral and multispectral satellite data are the two most commonly used datasets to identify natural and semi-natural vegetation. However, there is no documented analysis based on data from several areas concerning the difference in the classification accuracy of non-forest Natura 2000 habitat with the use of aerial hyperspectral and satellite multispectral data. Also, there is no recommendation, on which habitat can be classified with sufficient accuracy using free multispectral images. This study aimed to analyse the difference in classification accuracy of Natura 2000 habitats representing: meadows, grasslands, heaths and mires between data with different spectral resolutions and the results utility for nature conservation compared to conventional maps. The analysis was conducted in five study areas in Poland. The classification was performed on multispectral Sentinel-2 (S2) and hyperspectral HySpex (HS) images using the Random Forest algorithm. Based on the results, it can be stated that the use of HS data resulted in higher classification accuracy, on average 0.14, than using S2 images, regardless of the area of the habitat. However, the difference in accuracy was not constant, varying by area and habitat characterisation. Greater differences in accuracy were observed for areas where habitats were characterised by high α-diversity or ß-diversity. The HS and S2 data make it possible to create maps that provide a great deal of new knowledge about the distribution of Natura 2000 habitats, which is necessary for the management of protected areas. The obtained results indicate that by using S2 images it is possible to identify, at a satisfactory level, alluvial meadows and grassland. For heaths and mires, using HS data improved the results, but it is also possible to acquire general distribution of these classes, whereas HS images are obligatory for mapping salt, Molinia and lowland hay meadows.

Growth Stage-, Organ- and Time-Dependent Salt Tolerance of Halophyte Tripolium pannonicum (Jacq.) Dobrocz.

Tripolium pannonicum (Jacq.) Dobrocz. is a member of the diverse group of halophytes with the potential for the desalination and reclamation of degraded land. The adaptive processes of T. pannonicum to salinity habitats are still not well recognized. Therefore, we evaluated the effect of NaCl (0, 200, 400, and 800 mM) on: (1) two plant growth stages, (2) the activity of antioxidant enzymes and concentration of H2O2 and the proline in roots, stems, and leaves, and (3) the effect of long- and short-term salt stress on physiological responses. Germination, pot experiments, and a biochemical analysis were performed. The effective T. pannonicum's seed germination was achieved in the control. We demonstrated that halophyte's organs do not simply tolerate high-salt conditions. The activities of APX, POD, and catalase observed at 400 mM and 800 mM NaCl were varied between organs and revealed the following pattern: root > leaves > stem. Proline was preferentially accumulated in leaves that were more salt-tolerant than other organs. Salt stress enhanced the activity of antioxidant enzymes and concentrations of salinity stress indicators in a time-dependent manner. Our study has indicated that salt tolerance is a complex mechanism that depends on the growth phase, organs, and duration of salinity exposure. The results have potential for further proteomic and metabolomic analyses of adaptive salt tolerance processes.

Changes in physio-biochemical parameters and expression of metallothioneins in Avena sativa L. in response to drought.

Drought is one of the major threats to food security. Among several mechanisms involved in plant stress tolerance, one protein family-the plant metallothioneins (MTs)-shows great promise for enhancing drought resistance. Plant metallothioneins in oat (Avena sativa L.) have not yet been deeply analysed, and the literature lacks a comprehensive study of the whole family of plant MTs in response to drought. In this study, we showed that the number and nature of cis-elements linked with stress response in promoters of AsMTs1-3 differed depending on the MT type. Drought stress in oat plants caused an increase in the expression of AsMT2 and AsMT3 and a decrease in the expression of AsMT1 compared to well-watered plants. Moreover, the low values of relative water content, water use efficiency, net photosynthesis (PN), transpiration (E), stomatal conductance (gs), chlorophyll a, and carotenoid were accompanied by high levels of electrolyte leakage, internal CO2 concentration (Ci) and abscisic acid content, and high activity of antioxidants enzymes in plants under drought stress. The present study puts forward the idea that AsMTs are crucial for oat response to drought stress not only by regulating antioxidant activity but also by changing the plant water regime and photosynthesis. Our results support the hypothesis that structural differences among types of plant MTs reflect their diversified physiological roles.

Vegetation of temperate inland salt-marshes reflects local environmental conditions.

Inland salt marshes are recognized as habitats of unique and valuable vegetation at the European scale. There is still a lack of generalization regarding its vegetation syntaxonomy and environmental requirements, which is needed for its effective protection. To falsify our hypothesis about vegetation dependence on environmental requirements we aimed at description of the syntaxonomical units present in temperate European inland salt marshes and identification of their main environmental drivers. In our work we focused on the vegetation from the northern part of temperate salt marshes to limit confusion related to the geographical ranges of species. We collected the database of 968 vegetation plots from different European countries and applied the Cocktail method to analyze the data. Based on results, expert knowledge, existing syntaxonomical classifications and information from the literature, we identified diagnostic, constant and dominant species for individual syntaxonomical units. Then, we compiled maps of the vegetation unit distribution, and identified the most important environmental factors for the analyzed vegetation using statistical and multivariate methods, including canonical variate analysis. We classified the analyzed vegetation into nine classes, including two typical for salt-marsh vegetation - the Therosalicornietea and Festuco-Puccinellietea. Within these two classes, we distinguished two alliances and a total of five associations. The classes differ the most in terms of species preferences to salinity, soil moisture, light availability and soil nitrogen content. In addition salt marsh associations differ also by soil reaction and soil organic matter content. This provides direct implications for salt marsh sustainable management.

Understanding salinity stress responses in sorghum: exploring genotype variability and salt tolerance mechanisms.

Salinity, a significant abiotic stressor, adversely affects global plant growth. To address this, monitoring genetic diversity within a plant species germplasm for salt tolerance traits is vital. This study investigates the responses of ten sorghum genotypes to varying salt stress levels (control, 60 mM NaCl, and 120 mM NaCl), aiming to assess genetic diversity. Using a randomized complete block design with three replications and a split-plot arrangement, salt treatments were assigned to main plots, and genotypes were placed in sub-plots. Physiological attributes, including photosynthetic rate, stomatal conductance, CO2 concentration, leaf area index, chlorophyll concentrations, and antioxidant enzyme activity, were measured during the 50% flowering stage. Fresh forage yield was evaluated at the early dough stage, while dry forage yield and sodium/potassium concentrations were determined post-drying. Salinity induced 10-23% and 21-47% reductions in forage fresh yield at 60 mM and 120 mM NaCl, respectively, across sorghum genotypes. Forage dry yield also declined by 11-33% at 60 mM NaCl and 30-58% at 120 mM NaCl. Increased oxidative stress markers, proline, soluble carbohydrates, and antioxidant enzyme activity accompanied salinity. Genotypes exhibited diverse responses, with Payam showing significant chlorophyll and yield reductions at 60 mM NaCl and notable stress indicators at 120 mM NaCl. Pegah and GS4 demonstrated robust osmoregulation. In stress tolerance indices, Sepideh excelled at 60 mM NaCl, while GS4 outperformed at 120 mM NaCl. Pegah demonstrated high tolerance at 120 mM NaCl. Our findings highlight the importance of combating oxidative stress, managing water-related stress, and maintaining ionic homeostasis for sorghum's salt stress resilience. Key indicators like K/Na ratio, MDA, MSI, SOD, and proline effectively differentiate between tolerant and sensitive genotypes, offering valuable insights for sorghum breeding. Salt-tolerant sorghum genotypes exhibit stable photosynthesis, improved stomatal function, and membrane integrity through efficient osmotic regulation and robust antioxidant enzyme activity. This capability enables them to sustain performance, minimizing final product loss. The results suggest cultivating salt-tolerant sorghum in saline areas for increased sustainable production, with Pegah and GS4 emerging as promising candidates for further testing in salt-affected environments to obtain reliable yield data.

Salicornia europaea L. Functional Traits Indicate Its Optimum Growth.

Salicornia europaea L. grows in areas periodically flooded by salty or brackish water. It has potential economic value, because it can be used as food, forage, or biofuel, and has potential in pharmaceuticals and cosmetics. Increasing interest in S. europaea is due to its extreme salt tolerance and well growth in marginal saline soils. However, the variation in its functional traits in response to environmental conditions is still poorly studied. There are still questions regarding the optimal level of salinity for different traits. Therefore, we worked to address the question if S. europaea traits from different scales are controlled by salinity level. Based on performed pot experiment, we found that almost all traits are salinity dependent but affected in different ways. We demonstrated that morphological, biomass, and anatomical properties indicate optimum growth between 200 and 400 mM NaCl and growth limitations at 0, 800, and 1000 mM NaCl. Moreover, we found the most affected traits which include photosynthetic pigments and protein content, plant surface area, peroxidase activity, and anatomic traits related to cell shape. Our results significantly expanded the knowledge about S. europaea functional traits variation in response to salinity, which can be important for discovering regulating processes and for possible future agricultural applications.

Pseudomonas stutzeri and Kushneria marisflavi Alleviate Salinity Stress-Associated Damages in Barley, Lettuce, and Sunflower.

Soil salinity is one of the most important abiotic factors limiting plant productivity. The aim of this study was to determine the effect of selected halotolerant plant growth-promoting endophytes (PGPEs, Pseudomonas stutzeri ISE12 and Kushneria marisflavi CSE9) on the growth parameters of barley (Hordeum vulgare), lettuce (Lactuca sativa), and sunflower (Helianthus annuus) cultivated under salt stress conditions. A negative effect of two higher tested salinities (150 and 300 mM NaCl) was observed on the growth parameters of all investigated plants, including germination percentage and index (decreasing compared to the non-saline control variant in the ranges 5.3-91.7 and 13.6-90.9%, respectively), number of leaves (2.2-39.2%), fresh weight (24.2-81.6%); however, differences in salt stress tolerance among the investigated crops were observed (H. annuus > H. vulgare > L. sativa). Our data showed that the most crucial traits affected by endophyte inoculation under salt stress were chlorophyll concentration, leaf development, water storage, root development, and biomass accumulation. Thus, the influence of endophytes was species specific. K. marisflavi CSE9 promoted the growth of all tested plant species and could be considered a universal PGPEs for many plant genotypes cultivated under saline conditions (e.g., increasing of fresh weight compared to the non-inoculated control variant of barley, lettuce, and sunflower in the ranges 11.4-246.8, 118.9-201.2, and 16.4-77.7%, respectively). P. stutzeri ISE12 stimulated growth and mitigated salinity stress only in the case of barley. Bioaugmentation of crops with halotolerant bacterial strains can alleviate salt stress and promote plant growth; however, the selection of compatible strains and the verification of universal plant stress indicators are the key factors.

Foliar Application of Salicylic Acid Improves Salt Tolerance of Sorghum (Sorghum bicolor (L.) Moench).

It has been reported that around the world, approximately 19.5% of all irrigated land and 2.1% of dry land is affected by salt stress, and these percentages continue to increase. Sorghum is the fifth most important cereal in the world and therefore research on its salt tolerance is of global importance. In our research, we focused on foliar application of salicylic acid (SA) on salt-stressed sorghum. We performed a pot experiment with two salt levels (0 and 100 mM sodium chloride NaCl) and five SA concentrations (0, 50, 100, 150 and 200 mg/L). Our results suggest that in saline conditions foliar application of SA induced an adaptive response to salinity by inducing proline accumulation as well as antioxidant enzymes activities and enhanced the protection of the photosynthetic machinery, maintained photosynthesis activities, and improved the growth of sorghum plants. These alleviation effects were depended on applied SA concentration. Under saline condition 150 mg/L, SA was the most effective for relieving the adverse effect of salt stress. Under non-saline conditions 100 mg/L SA was the best for improving sorghum growth and dry matter production. Our results demonstrated that foliar SA application is effective in improving sorghum growth under salinity.

Invasive Plant Species Distribution Is Structured by Soil and Habitat Type in the City Landscape.

Invasive alien species (IAS) is a global problem that largely relates to human activities and human settlements. To prevent the further spread of IAS, we first need to know their pattern of distribution, to determine which constitutes the greatest threat, and understand which habitats and migration pathways they prefer. Our research aimed to identify the main vectors and distribution pattern of IAS of plants in the city environment. We checked the relations between species distribution and such environmental factors as urban soil type and habitat type. We applied data on IAS occurrence (collected in the period 1973-2015) in 515 permanent plots with dimensions of 0.5 × 0.5 km and analyzed by direct ordination methods. In total, we recorded 66 IAS. We found a 27% variance in the IAS distribution pattern, which can be explained by statistically significant soil and habitat types. The most important for species distribution were: river and alluvial soils, forests and related rusty soils, and places of intensive human activities, including areas of urbisols and industriosols. Our results provide details that can inform local efforts for the management and control of invasive species, and they provide evidence of the different associations between natural patterns and human land use.

How Does Salinity Shape Bacterial and Fungal Microbiomes of Alnus glutinosa Roots?

Black alder (Alnus glutinosa Gaertn.) belongs to dual mycorrhizal trees, forming ectomycorrhizal (EM) and arbuscular (AM) root structures, as well as represents actinorrhizal plants that associate with nitrogen-fixing actinomycete Frankia sp. We hypothesized that the unique ternary structure of symbionts can influence community structure of other plant-associated microorganisms (bacterial and fungal endophytes), particularly under seasonally changing salinity in A. glutinosa roots. In our study we analyzed black alder root bacterial and fungal microbiome present at two forest test sites (saline and non-saline) in two different seasons (spring and fall). The dominant type of root microsymbionts of alder were ectomycorrhizal fungi, whose distribution depended on site (salinity): Tomentella, Lactarius, and Phialocephala were more abundant at the saline site. Mortierella and Naucoria (representatives of saprotrophs or endophytes) displayed the opposite tendency. Arbuscular mycorrhizal fungi belonged to Glomeromycota (orders Paraglomales and Glomales), however, they represented less than 1% of all identified fungi. Bacterial community structure depended on test site but not on season. Sequences affiliated with Rhodanobacter, Granulicella, and Sphingomonas dominated at the saline site, while Bradyrhizobium and Rhizobium were more abundant at the non-saline site. Moreover, genus Frankia was observed only at the saline site. In conclusion, bacterial and fungal community structure of alder root microsymbionts and endophytes depends on five soil chemical parameters: salinity, phosphorus, pH, saturation percentage (SP) as well as total organic carbon (TOC), and seasonality does not appear to be an important factor shaping microbial communities. Ectomycorrhizal fungi are the most abundant symbionts of mature alders growing in saline soils. However, specific distribution of nitrogen-fixing Frankia (forming root nodules) and association of arbuscular fungi at early stages of plant development should be taken into account in further studies.

Ectomycorrhizal and endophytic fungi associated with Alnus glutinosa growing in a saline area of central Poland.

Alnus glutinosa (black alder) is a mycorrhizal pioneer tree species with tolerance to high concentrations of salt in the soil and can therefore be considered to be an important tree for the regeneration of forests areas devastated by excessive salt. However, there is still a lack of information about the ectomycorrhizal fungi (EMF) associated with mature individuals of A. glutinosa growing in natural saline conditions. The main objective of this study was to test the effect of soil salinity and other physicochemical parameters on root tips colonized by EMF, as well as on the species richness and diversity of an EMF community associated with A. glutinosa growing in natural conditions. We identified a significant effect of soil salinity (expressed as electrical conductivity: ECe and EC1:5) on fungal taxa but not on the total level of EM fungal colonization on roots. Increasing soil salinity promoted dark-coloured EMF belonging to the order Thelephorales (Tomentella sp. and Thelephora sp.). These fungi are also commonly found in soils polluted with heavy-metal. The ability of these fungi to grow in contaminated soil may be due to the presence of melanine, a natural dark pigment and common wall component of the Thelephoraceae that is known to act as a protective interface between fungal metabolism and biotic and abiotic environmental stressors. Moreover, increased colonization of fungi belonging to the class of Leotiomycetes and Sordiomycetes, known as endophytic fungal species, was observed at the test sites, that contained a larger content of total phosphorus. This observation confirms the ability of commonly known endophytic fungi to form ectomycorrhizal structures on the roots of A. glutinosa under saline stress conditions.

Ectomycorrhizal Community Structure of Salix and Betula spp. at a Saline Site in Central Poland in Relation to the Seasons and Soil Parameters.

Saline stress is one of the most important abiotic factors limiting the growth and development of plants and associated microorganisms. While the impact of salinity on associations of arbuscular fungi is relatively well understood, knowledge of the ectomycorrhizal (EM) fungi of trees growing on saline land is limited. The main objective of this study was to determine the density and diversity of EM fungi associated with three tree species, Salix alba, Salix caprea and Betula pendula, growing in saline soil during two seasons, autumn and spring. The site was located in central Poland, and the increased salinity of the soil was of anthropogenic origin from soda production. The degree of EM colonisation of fine root tips varied between 9 and 34 % and depended on the tree species of interest (S. caprea < S. alba < B. pendula) and season (spring < autumn). Moreover, the ectomycorrhizal colonisation of B. pendula was positively correlated with pH and CaCO3, while for S. caprea and S. alba, colonisation was associated with most of the other soil parameters investigated; e.g. salinity, Corg and N. Analysis of EM fungi revealed four to five different morphotypes per each season: Tomentella sp. Sa-A, Hebeloma collariatum Sc-A, Geopora sp. Sc-A, Helotiales sp. Bp-A in the autumn and Tomentella sp. Sa-S, Tomentella sp. Sc-S and three morphotypes from the families Thelephoraceae and Pyronemataceae in the spring. In conclusion, the density of EM is related to the level of salinity (ECe), season and tree species. Tomentella spp., Hebeloma sp., Geopora sp. and Helotiales sp. are groups of species highly adapted to saline conditions.