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Laboratory experimentsCulturingThe marine cyanobacterium Trichodesmium erythraeum IMS101 was obtained from the National Center for Marine Algae and Microbiota (Maine, USA) and was grown in Aquil-tricho medium prepared with 0.22 µm-filtered and microwave-sterilized oligotrophic South China Sea surface water6. The medium was enriched with various concentrations of chelexed and filter-sterilized NaH2PO4 as where indicated, and filter-sterilized vitamins and trace metals buffered with 20 µM EDTA6. The cultures were unialgal, and although they were not axenic, sterile trace metal clean techniques were applied for culturing and experimental manipulations. T. erythraeum was pre-adapted to low P condition by semi-continuously culturing at 0.5 μM PO43− and at two pCO2 levels (400 and 750 µatm) for more than one year. To start the chemostat culture, three replicates per treatment were grown in 1-L Nalgene® magnetic culture vessels (Nalgene Nunc International, Rochester, NY, USA), in which the cultures were continuously mixed by bubbling with humidified and 0.22 µm-filtered CO2–air mixtures and stirring using a suspended magnetic stir bar. The reservoirs contained Aquil-tricho medium with 1.2 μM NaH2PO4, which was delivered to the culture vessels using a peristaltic pump (Masterflex® L/S®, USA) at the dilution rate of 0.2 d−1. In all experiments, cultures were grown at ;27 °C and ~80 μmol photons m−2 s−1 (14 h:10 h light–dark cycle) in an AL-41L4 algae chamber (Percival). The concentration of Chlorophyll a (Chla) was monitored daily in the middle of the photoperiod as an indicator of biomass. When the Chla concentration remained constant for more than one generation, the system was considered to have reached steady-state, and was maintained for at least another four generations prior to sampling for further analysis.Carbonate chemistry manipulationpCO2/pH of seawater media in the culture vessels and in the reservoir was controlled by continuously bubbling with humidified and 0.22 µm-filtered CO2-air mixtures generated by CO2 mixers (Ruihua Instrument & Equipment Ltd.). During the experimental period, the pHT (pH on the total scale) of media was monitored daily using a spectrophotometric method46. The dissolved inorganic carbon (DIC) of media was analyzed by acidification and subsequent quantification of released CO2 with a CO2 analyzer (LI 7000, Apollo SciTech). Calculations of alkalinity and pCO2 were made using the CO2Sys program47, based on measurements of pHT and DIC, and the carbonate chemistry of the experiments are shown in Supplementary Table 1.Chla concentration and cell density and sizeChla concentration was measured daily following Hong et al.6. Briefly, T. erythraeum was filtered onto 3 μm polycarbonate membrane filters (Millipore), followed by heating at 65 °C for 6 min in 90% (vol/vol) methanol. After extraction the filter was removed and cell debris were spun down via centrifugation (5 min at 20,000×g) before spectrophotometric analysis. Cell density and the average cell length and width were determined at regular intervals when the chemostat cultures reached steady-state using ImageJ software. Photographs of Trichodesmium were taken using a camera (Canon DS126281, Japan) connected with an inverted microscope (Olympus CKX41, Japan). Total number and length of filaments in 1 mL of culture were measured, and the cell number of ~20 filaments was counted. The average length of cells was obtained by dividing the total length of the 20 filaments by their total cell number. The cell density of the culture was then calculated by dividing the total length of filaments in 1 mL culture by the average cell length. The average cell width was determined by measuring the width of around 1000 cells in each treatment.Elemental compositionTo determine particulate organic C (POC) and N (PON), at the end of the chemostat culturing T. erythraeum cells were collected on pre-combusted 25 mm GF/F filters (Whatman) and stored at −80 °C. Prior to analysis, the filters were dried overnight at 60 °C, treated with fuming HCl for 6 h to remove all inorganic carbon, and dried overnight again at 60 °C. After being packed in tin cups, the samples were subsequently analyzed on a PerkinElmer Series II CHNS/O Analyzer 2400.Particulate organic P (POP) was measured following Solorzano et al.48. Cells were filtered on pre-combusted 25 mm GF/F filters and rinsed twice with 2 mL of 0.17 M Na2SO4. The filters were then placed in combusted glass bottles with the addition of 2 mL of 0.017 M MgSO4, and subsequently evaporated to dryness at 95 °C and baked at 450 °C for 2 h. After cooling, 5 mL of 0.2 M HCl was added to each bottle. The bottle was then tightly capped and heated at 80 °C for 30 min, after which 5 mL Milli-Q H2O was added. Dissolved phosphate from the digested POP sample was measured colorimetrically following the standard phosphomolybdenum blue method.C uptake and N2 fixation ratesRates of short-term C uptake were determined at the end of the chemostat culturing. 100 µM NaH14CO3 (PerkinElmer) was added to 50 mL of cultures in the middle of the photoperiod, which was then incubated for 20 min under the growth conditions. After incubation, the samples were collected onto 3 μm polycarbonate membrane filters (Millipore), which were then washed with 0.22 µm-filtered oligotrophic seawater and placed on the bottom of scintillation vials. The filters were acidified to remove inorganic C by adding 500 µL of 2% HCl. The radioactivity was determined using a Tri-Carb 2800TR Liquid Scintillation Analyzer (PerkinElmer). Rates of N2 fixation (nitrogenase activity) were measured in the middle of the photoperiod for 2 h by the acetylene reduction assay49, using a ratio of 4:1 to convert ethylene production to N2 fixation.Soluble reactive phosphate (SRP) analysisWhen the chemostat cultures reached a steady-state, SRP concentrations in the culture vessels were measured at regular intervals, using the classic phosphomolybdenum blue (PMB) method with an additional step to enrich PMB on an Oasis HLB cartridge50. Briefly, 100 mL of GF/F filtered medium sample was fortified with 2 mL of ascorbic acid (100 g L−1) and 2 mL of mixed reagent (MR, the mixture of 100 mL of 130 g L−1 ammonium molybdate tetrahydrate, 100 mL of 3.5 g L−1 potassium antimony tartrate, and 300 mL of 1:1 diluted H2SO4), and then mixed completely. After standing at room temperature for 5 min, the solution was loaded onto a preconditioned Oasis HLB cartridge (3 cm3/60 mg, P/N: WAT094226, Waters Corp.) via a peristaltic pump, and then 1 mL eluent solution (0.2 M NaOH) was added to elute the sample into a cuvette, to which 0.06 mL of MR and 0.03 mL of ascorbic acid solution was added to fully develop PMB. Finally, the absorbance of PMB was measured at 700 nm using a spectrophotometer.Alkaline phosphatase (AP) activityAP activities were measured in the middle of the photoperiod using p-nitrophenylphosphate (pNPP) as a substrate51. Briefly, 5 mL of culture was incubated with 250 μL of 10 mM pNPP, 675 μL of Tris-glycine buffer (50 mM, pH 8.5) and 67.5 μL of 1 mM MgCl2 for 2 h under growth conditions. The absorbance of formed p-nitrophenol (pNP) was measured at 410 nm using a spectrophotometer.PolyP analysisAt the end of the chemostat culturing, T. erythraeum cells were filtered in the middle of the photoperiod onto 3 μm polycarbonate membrane filters (Millipore), flash frozen in liquid nitrogen, and stored at −80 °C until analysis. PolyP was quantified fluorometrically following Martin and Van Mooy22 and Martin et al.23. Briefly, samples were re-suspended in 1 mL Tris buffer (pH 7.0), sonicated for 30 s, immersed in boiling water for 5 min, sonicated for another 30 s, and then digested by 10 U DNase (Takara), RNase (2.5 U RNase A + 100 U RNase T1) (Invitrogen) and 20 μl of 20 mg mL−1 proteinase K at 37 °C for 30 min. After centrifugation for 5 min at 14,000×g, the supernatant was diluted with Tris buffer according to the range of standards curve, stained with 60 μL of 100 μM 4, 6-diamidino-2-phenylindole (DAPI) per 500 μL of samples, incubated for 7 min and then vortexed. The samples were then loaded onto a black 96-well plate and the absorption of fluorescence at an excitation wavelength of 415 nm and emission wavelength of 550 nm was measured using a PerkinElmer EnSpire® Multimode Plate Reader. PolyP standard (sodium phosphate glass Type 45) was purchased from Sigma-Aldrich. This method gives a relative measure of polyP concentration23 that is expressed as femto-equivalents of the standard per cell (feq cell−1).Cellular ATP measurementCellular ATP contents were determined when the chemostat cultures reached a steady state. T. erythraeum cells were collected in the middle of the photoperiod using an ATP Assay Kit (Beyotime Biotechnology, Shanghai, China) according to the manufacturer’s instructions. Briefly, the sample was lysed and centrifuged, and the supernatant (100 μL) was mixed with ATP detection working reagent (100 μL) and loaded onto a black 96-well plate. The luminescence was measured using a PerkinElmer EnSpire® Multimode Plate Reader.Intracellular metabolites measurementsNAD(H), NADP(H), and Glu were measured at the end of the chemostat culturing, using the liquid chromatography-tandem quadrupole mass spectrometry (LC–MS/MS) method modified from Luo et al.52. Briefly, T. erythraeum cells were gently filtered at the middle of photoperiod onto 3 μm polycarbonate membrane filters (Millipore), rapidly suspended in −80 °C precooled methanol-water (60%, v/v) mixture. After being kept in −80 °C freezer for 30 min, the sample was sonicated for 30 s, centrifuged at 12,000×g and 4 °C for 5 min, and the supernatant was filtered through a 0.2 μm filter (Jinteng®, China) and stored at −80 °C for further LC–MS/MS analysis.A 2.0 × 50 mm Phenomenex® Gemini 5u C18 110 Å column (particle size 5.2 µm, Phenomenex, USA) was used for the analysis. The mobile phases consisted of two solvents: mobile phase A (10 mM tributylamine aqueous solution, pH 4.95 with 15 mM acetic acid) and mobile phase B (100% methanol), which were delivered using an Agilent 1290 UPLC binary pump (Agilent Technologies, Palo Alto, CA, USA) at a flow rate of 200 µL min−1, with a linear gradient program implemented as follows: hold isocratic at 0% B (0–2 min); linear gradient from 0% to 85% B (2–28 min); hold isocratic at 0% B (28–34 min). The effluent from the LC column was delivered to an Agilent 6490 triple-quadrupole mass spectrometer, equipped with an electrospray ionization source operating in negative-ion mode. NAD, NADH, NADP, NADPH, and Glu were monitored in the multiple reaction monitoring modes with the transition events at m/z 662.3  > 540, 664.3  > 79, 742  > 620, 744  > 79, and 147  > 84, respectively.RNA extraction, library preparation, and sequencingAt the end of the chemostat culturing, T. erythraeum was collected in the middle of the photoperiod by filtering onto 3 μm polycarbonate membrane filters (Millipore), flash frozen in liquid nitrogen and stored at −80 °C until extraction. Total RNA was extracted using TRIzol® Reagent (Invitrogen) combined with a physical cell disruption approach by glass beads according to the manufacturer’s instructions. Genomic DNA was removed thoroughly by treating it with RNAase-free DNase I (Takara, Japan). Ribosomal RNA was removed from a total amount of 3 µg RNA using Ribo-Zero rRNA Removal kit (Illumina, USA). Subsequently, cDNA libraries were generated according to the manufacturer’s protocol of NEBNext® UltraTM Directional RNA Library Prep Kit for Illumina® (NEB, USA). The quality of the library was assessed on the Agilent Bioanalyzer 2100 system (Agilent Technologies, CA, USA). Libraries were sequenced on an Illumina Hiseq 2500 platform, yielding 136-bp paired-end reads.RNA-Seq bioinformaticsClean reads were obtained from raw data by removing reads containing adapter, ploy-N and low-quality read. Qualified sequences were mapped to the Trichodesmium erythraeum IMS101 genome (https://www.ncbi.nlm.nih.gov/nuccore/NC_008312.1) by using Bowtie2-2.2.353. Differential expression analysis for high/low pCO2 with P limitation was performed using the DESeq2 R package54. The resulting p-values were adjusted using Benjamini and Hochberg’s approach for controlling the false discovery rate. Genes with an adjusted p-value  More

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Soil sampling and determination of soil physical properties and synoptic dataSoil samples were taken from two coastal deserts in the north and south of Iran. Details of their geographic distribution and eco-physiological characterization were shown in Table 1. A total of 2 kg of soil samples were collected from 2 distinct sampling locations ranging in depth from 0 to 30 cm, and the samples were dried for 3 days at room temperature and in the dark before sifting. The soil samples were sieved using a 2 mm sieve to remove stones and other inert material before being stored in zip-top bags. Table 1 lists the soil samples’ physical characteristics, including soil texture (sand 2–0.02 mm; silt 0.02–0.002 mm; clay 0.002 mm), pH, and the proportions of clay, silt, and sand. Synoptic data from the past 10 years (2009–2019), including the average annual temperature, maximum temperature, minimum temperature, average rainfall, average annual wind speed, and maximum wind speed, were obtained from the I.R.OF Iran Meteor (http://www.irimo.ir/far/index.php).Bacterial isolation and effect of manure-based medium on their growthAccording to Chen et al. 2005, the soil-borne bacteria were isolated using direct-spreading method. For this essence soil samples were treated through a series of dilutions. The mixture of 1 g of soil sample was vortexed for 1 min after being suspended in 2 ml of sterile physiological saline (0.9% w/v NaCl). The mixture was then diluted serially (typically 10–1 to 10–7), and level 100 μl of the diluted soil samples were scattered on the surface of solidified plates using glass spreaders. The samples were then incubated for 1 to 3 days at 30 °C in an inverted posture without light. For bacterial isolation, we used eleven culture media including Nutrient Agar (NA), Nutrient Agar plus MnSO4 (NA + MnSO4), LB, Moller Hinton Agar (MHA), Acidithiobacillus (APH) medium, Violet Red Bile Lactose (VRB) agar medium, GYM Streptomyces medium, DPM medium, Azospirillum medium, Azotobacter medium and Manure based medium (MB).To prepare MB medium, dry animal manure and distilled water (1:6 w/v) were combined to create MB medium, which was then let to sit at room temperature for 16 h. The resulting mixture was then centrifuged at 5000 rcf for 30 min after being filtered twice. The next stage involved adding Hoagland salts (10% w/v) to the final extract, adjusting the medium’s pH to 5.8 ± 0.02, and autoclaving it for 20 min at 121 °C and 1.5 kPa. Before sterilization, bacteriological agar (1.5 w/v) was employed as a gelling agent to solidify the medium.After bacterial isolation on NA, NA+ MnSO4, LB, MHA, APH, VRB, GYM, DPM, and Azospibrillum media, the growth of all isolates was evaluated on an MB medium. To investigate isolates biomass in the same condition, we elected MB medium. First, the bacteria were grown in the liquid form of NA, NA+ MnSO4, LB, MHA, APH, VRB, GYM, DPM, and Azospirillum and Azotobacter media at 30 °C for 48 h, then 103 cells of each isolate were transferred to 48 wells plates containing MB medium, and plates were incubated at 30 °C for 10 h. Then, the growth of bacteria was read at an optical density (OD) of 630 nm 10 h after inoculation, the experiment was performed with three replicates. In the following step, CFU/ml equivalent to each OD was obtained by inoculating the uniform amount of liquid culture of the isolates on the solid form of MB medium at 30 °C for 16 h.Phenotypic characterization and biochemical identification of bacterial isolatesThe morphological analysis of the cell shape, colony (i.e., shape, color, and size), and biochemical tests were used to identify the bacterial isolates. Biochemical characterization was carried out By using gram staining, KOH27, oxidase, and catalase tests. For this essence, following Bartholomew’s method28, gram staining of bacteria was studied 48 h after inoculation on MHA, and the non-staining KOH method was used to confirm the results. Using 0.5 ml of a 10% hydrogen peroxide solution, a catalase test was conducted, and the generation of gas bubbles was monitored. Using biochemical oxidase discs, the oxidative activity of 27 isolates was investigated.Effect of abiotic stresses on bacterial isolatesTo determine the effect of abiotic stresses on isolates alkaline (MH medium with pH  10), salinity (MH medium supplemented with the final concentration of 100 mM NaCl), osmotic [MH medium supplemented with 25% polyethylene glycol (PEG) Mn6000], and thermal stresses (MH medium incubated at 15 °C for cold stress and 60 °C for heat stress) were screened. For all experiments, the incubation period was 15 h, and plates were kept in a dark condition.MALDI-TOF MS identification of isolatesSoil bacterial isolates were subcultured twice on MHA and incubated at 30 °C for 24 h before MALDI-TOF MS measurement. Then ∼0.1 µg of cell material was directly transferred from a bacterial colony or smear of colonies to a MALDI target spot. After drying at laboratory temperature, sample spots were overlaid with 1 μl of matrix solution (10 mg/mL a-cyano-4-hydroxycinnamic acid in 50% acetonitrile and 2.5% trifluoroacetic acid) and each measurement was carried out in triplicate (technical replicates). MS analysis was performed on an Autoflex MALDI-TOF mass spectrometer (Bruker Daltonics, Germany) using Flex Control 3.4 software (Bruker Daltonics, Germany). Calibration was carried out with the use of the Bacterial Test Standard (Bruker Daltonics, Germany). Soil isolates with a valid MALDI-TOF MS score of 2 were undoubtedly assigned to the genus/species level. For bacterial classification and identification, BioTyper 3.1 software (Bruker Daltonics, Germany) equipped with MBT 6903 MPS Library (released in April 2016), the MALDI Biotyper Preprocessing Standard Method, and the MALDI Biotyper MSP Identification Standard Method adjusted by the manufacturer (Bruker Daltonics, Germany) were used. Only the highest score value of all mass spectra belonging to individual cultures (biological and technical replicates) was recorded25. The score between 2.3 and 3.00 shows highly probable species-level identification and between 2.0 and 2.29 represents genus-level identification and probable species level of identification. A score between 1.7 and 1.99 indicates probable genus-level identification29.Effects of bacterial isolates on plants growthThe Seed and Plant Improvement Institute of Karaj (Karaj, Iran; http://www.spii.ir/homepage.aspx?site=DouranPortal&tabid=1&lang=faIR) provided the maize, canola, and wheat seeds (Zea mays. Var Kosha; Brassica napus Var Nima; Triticum aestivum Var Kalate). In greenhouse trials, 2 × 103 cells/seed of soil-borne isolates cultured in a manure-based medium were inoculated to maize, canola, and wheat plants. During the studies, sand that had been acid washed and autoclaved was used for planting. For three weeks, seedlings were kept under a 16/8 h day/night photoperiod with a 25 °C temperature. Three replications of a complete randomized block design were used for the colonization experiment’s treatments. Under the bacterial treatments, measurements were made of the plant growth parameters including shoot dry biomass (mg), root dry biomass (mg), shoot length (cm), root length (cm), shoot density (mg/cm), root density (mg/cm), and shoot/root weight (mg). Samples were dried at 60 °C for three days to measure dry biomass.Statistical analysisStatistical analysis was done by R software (version 4.1.3). One-way analysis of variance (ANOVA) was used to determine the significance of the experiment, and Fisher’s protected Least Significant Difference (LSD) test with a P-value of 0.01 was performed to separate the means. Furthermore, PCA analysis has been carried out based on the Clustvis package and the SVD imputation approach.Ethics approval and consent to participateAll authors agree to the ethics and consent to participate in this article and declare that this submission follows the policies of Scientific Reports. Accordingly, the material is the author’s original work, which has not been previously published elsewhere. The paper is not being considered for publication elsewhere. All authors have been personally and actively involved in substantial work leading to the paper and will take public responsibility for its content.Ethics for research involving plantsAll authors confirmed that experimental research and field studies on plants, including receiving the seeds from the Seed and Plant Improvement Institute of Karaj, complied with relevant institutional, national, and international guidelines and legislation. Furthermore, methods were conducted according to the relevant guidelines and regulations. More