Oxygen-mycorrhiza synergy drives phosphorus transformation in soil and its accumulation in greenhouse-grown tomato.

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Yingji Lian, Hongjun Lei, Hongwei Pan, Muhammad Zain, Xin Liu, and 3 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7489363/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Background and Aims: Limited soil phosphorus (P) availability impairs plant growth in protected cultivation systems. Although arbuscular mycorrhizal fungi (AMF) can enhance P uptake, the combined effects of AMF and aerated drip irrigation (ADI) on soil P fractions and tomato P accumulation remain unclear. Methods We conducted a greenhouse experiment with two irrigation regimes—standard drip (DO = 6 mg·L⁻¹) and ADI (DO = 15 mg·L⁻¹)—and four AMF treatments: control, Funneliformis mosseae (FM), Rhizophagus intraradices (RI), and FM + RI mixed inoculation. We assessed AMF colonization, soil P fractions, tomato P accumulation, yield, and employed structural equation modeling (SEM) to elucidate mechanisms. Results ADI combined with RI (ARI) significantly enhanced AMF colonization (+ 32.66%, P < 0.05), increased the readily available Resin-P fraction by 32.09%, promoted organic-to-inorganic P conversion (NaHCO 3 -Pi + 15.87%, NaHCO 3 -Po − 23.64%, NaOH-Po − 20.57%), and resulted in 34.4% greater plant P accumulation and 36.8% higher yield compared to RI under standard irrigation. SEM revealed two key mechanisms: increased acid phosphatase activity driving organic P mineralization and optimized root morphology enhancing P uptake. Conclusion Our findings support that ADI and AMF synergistically improve P availability, uptake, and tomato productivity by integrating soil biochemical transformation and enhanced root architecture—offering a promising strategy for sustainable phosphorus management in greenhouse production. Aerated drip irrigation Arbuscular mycorrhizal fungi Mycorrhizal colonization rate Phosphorus transformation Phosphorus accumulation Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 1. Introduction Phosphorus is an essential macronutrient for plant growth and development, playing a critical role in energy transfer, photosynthesis, nutrient transport, and metabolic processes. It is particularly important for root development, flowering, and fruit formation in tomato ( Solanum lycopersicum L. ), directly affecting both yield and quality (Khan et al. 2023). However, in controlled agricultural systems, due to high crop rotation indices and excessive fertilization, phosphorus in the soil is easily fixed by iron, aluminum oxides, or calcium salts, forming insoluble phosphates. As a result, the proportion of available phosphorus to plants is typically less than 20% (Zhang et al. 2019; Fang et al. 2023; Négrel et al. 2024). The low phosphorus use efficiency in controlled systems is a critical bottleneck limiting the high yield and sustainability of greenhouse tomato production. Traditional strategies rely on increasing the application of phosphorus fertilizers, which not only raises production costs but also leads to significant accumulation of residual phosphorus in the soil (Pavinato et al. 2020), which ultimately results in soil degradation and environmental issues. Therefore, there is an urgent need to explore innovative technologies that can effectively activate fixed phosphorus in the soil and improve plant phosphorus absorption and utilization efficiency (Song et al. 2020). Researchers have actively explored solutions to address the issue of phosphorus fixation. Arbuscular mycorrhizal fungi (AMF) are soil microorganisms closely related to plant nutrition (Tedersoo et al. 2020) that form symbiotic relationships with approximately 80% of terrestrial plants (Branco et al. 2022; Shi et al. 2023). These AMF are reported to be involved in significantly extending the root absorption range and enhancing the uptake of water and nutrients (Kuila and Ghosh 2022), particularly phosphorus (Peng et al. 2020). The hyphal network of AMF can directly absorb phosphates from the soil and transfer them to the plant roots through the mycorrhizal interface. Additionally, AMF can promote the mineralization of organic phosphorus by secreting acid phosphatases (Chen et al. 2020). Aerated drip irrigation (ADI), an advanced version of subsurface drip irrigation, improves the soil dissolved oxygen content, which enhances the rhizosphere microecological environment, increases soil oxygen levels, and the air-filled porosity (Ouyang et al. 2025; Wang et al. 2024). This in turn boosts microbial activity, promotes root growth, and facilitates nutrient transformation. Dong et al. (2025) found that ADI significantly increased root length by 19.02%, root volume by 22.92%, and root vitality by 32.1% as compared to traditional drip irrigation. Furthermore, ADI can indirectly promote nutrient cycling by altering soil microbial community structure and increasing the abundance of beneficial microorganisms (Zhao et al. 2019; Zhu et al. 2022). Research on vertical flow constructed wetlands has shown that intermittent aeration can increase the AMF colonization rate to 67% (Xu et al. 2021), indicating that moderate aeration can optimize AMF community structure. Although both ADI and AMF have shown progress in improving phosphorus use efficiency when applied individually, but still there is limited systematic research on their synergistic effects. The environmental factors such as high temperature, high humidity, and low oxygen levels may weaken AMF colonization efficiency and function, specifically in controlled soils (Carvalho et al. 2015; Huo et al. 2021). In addition, the application of ADI alone may not fully exert its regulatory potential on soil microbial communities, limiting its effectiveness in enhancing phosphorus utilization and plant stress resistance (Cao et al. 2024; Chen et al. 2023). The research gap hinders the development of strategies to optimize phosphorus management in controlled agricultural systems. Therefore, we hypothesized that comprehensive study on the synergistic effects of ADI and AMF on soil phosphorus transformation, root morphology, and phosphorus allocation, as well as the interactive effects of different AMF strains (such as Rhizophagus intraradices and Funneliformis mosseae ) under oxygenated conditions should be conducted. We planned a study with objectives to ( 1 ) investigates the synergistic effects of ADI and AMF through a two-factor experiment (irrigation mode × AMF inoculation) and systematically analyzes the changes in Hedley soil phosphorus fractions, root morphology, and soil phosphatase activity in response to oxygen-mycorrhiza synergy; ( 2 ) quantify the contribution of AMF colonization rate and soil phosphorus fraction changes to phosphorus uptake and yield in tomato plants through Structural equation modeling (SEM). The findings from this study will provide new insights into addressing phosphorus fixation in controlled soils and improving phosphorus fertilizer use efficiency. 2. Materials and methods 2.1 Experimental materials The tested tomato variety Solanum lycopersicum "Huafei No. 1", was acquired from the Xinxiang Breeding Base. Two types of arbuscular mycorrhizal fungi (AMF) inoculants were used: one was Funneliformis mosseae (FM), purchased from the China AMF Resource Bank (BGC, XJ01) at the Beijing Academy of Agriculture and Forestry Sciences. The spore concentration was approximately 230 spores g − 1 , including the corresponding culture medium, spores, mycelium, and plant root segments. The other inoculant Rhizophagus intraradices (RI) was taken from Foshan Ironman Environmental Technology Co., Ltd. The inoculant contained fine sand, spores, mycelium, and plant root segments, with a spore concentration of approximately 70 spores g − 1 . The experiment was conducted in pots filled with clay soil. The pots were placed in a modern greenhouse at the Agricultural Efficient Water Use Experimental Field of North China University of Water Resources and Electric Power. The greenhouse was equipped with wet curtains, fans, and shading nets to regulate temperature and humidity. The soil was sterilized by high-temperature treatment: it was first sieved through a 2 mm mesh, then placed in an oven set to 160°C for 2 hours. After cooling, the sterilization process was repeated for another 2 hours. The soil was left to cool naturally before use (Hu et al. 2020). The basic chemical properties of the soil are given in Table 1 . Table 1 Basic chemical properties of tested soils. Soil type Bulk density (g·cm − 3 ) Field capacity (%) Organic matter (g·kg − 1 ) Nitrate nitrogen (mg·kg − 1 ) Ammonium nitrogen (mg·kg − 1 ) Total phosphorus (g·kg − 1 ) Available phosphorus (mg·kg − 1 ) pH Clay 1.32 25.94 11.62 21.4 4.55 0.70 58.43 8.11 2.2 Experimental design The experiment was a two-factor fully factorial design with the following treatments. ( 1 ) Irrigation Modes: (a) Conventional standard drip irrigation without additional aeration (C) has a dissolved oxygen (DO) value of approximately 6 mg·L − 1 in the irrigation water; (b) Aerated drip irrigation (A) with a DO value set to 15 mg·L − 1 in the irrigation water; ( 2 ) AMF Inoculation Treatments: (a) Funneliformis mosseae (FM) inoculation only; (b) Rhizophagus intraradices (RI) inoculation only; (c) A mixed inoculation with both fungi, each at 50% of the individual spore concentration (FMRI). Based on the spore concentration of the inoculants, the FM treatment received 5g of Funneliformis mosseae , the RI treatment received 16g of Rhizophagus intraradices , and the FMRI treatment received 2.5g of Funneliformis mosseae and 8g of Rhizophagus intraradices . This resulted in a total of 8 treatments, with 5 replications per treatment, totaling 40 pots. The details of experimental design are presented in Table 2 . Table 2 Experimental treatments with their respective DO and inoculants concentrations. Treatment DO in irrigation water (mg·L − 1 ) Funneliformis mosseae (g·pot − 1 ) Rhizophagus intraradices (g·pot − 1 ) C 6 0 0 CRI 6 0 16 CFM 6 5.0 0 CFMRI 6 2.5 8 A 15 0 0 ARI 15 0 16 AFM 15 5.0 0 AFMRI 15 2.5 8 The numbers in the table represent the levels of different independent variables. DO represents the dissolved oxygen concentration of irrigation water; C represents standard drip irrigation without additional aeration; A represents aerated drip irrigation; RI refers to the treatment with Rhizophagus intraradices (RI) inoculation only; FM refers to the treatment with Funneliformis mosseae (FM) inoculation only; FMRI refers to the mixed inoculation of FM and RI. The pots used for the experiment were 22 cm in diameter and 27 cm in height. After sterilizing the soil with high-temperature treatment, each pot was filled with 8 kg of dry soil, mixed with 2.0 g of calcium superphosphate as a basal fertilizer. This mixture was irrigated with 500 mL of water on the same day, and the soil was left to settle and infiltrate overnight, followed by supplemental watering based on the soil moisture condition on the next day until the soil moisture content reached approximately 70% of the field capacity, at which point transplanting was conducted. A hole (3–5 cm deep) was dug in each pot, arbuscular mycorrhizal fungi (AMF) were placed in the hole in a flat layer, and then one tomato seedling with 4 true leaves and 1 heart or 5 true leaves and 1 heart was transplanted into the pot, followed by compaction of the surrounding soil. After transplantation, a soil solution was prepared by mixing the soil with deionized water at a 2:1 soil-to-water ratio, and the mixture was wet-sieved through a 30 µm filter to remove large soil microorganisms, including AMF spores. The filtrate was then quantified (20 ml·pot − 1 ) and added to the pots to restore the basic microbial community of the sterilized soil (Sun et al. 2022). Fertilization management : Each pot was fertilized with urea (46% N content), calcium superphosphate (12% P content), and potassium sulfate (50% K content) as nitrogen, phosphorus, and potassium fertilizers, respectively. Corresponding to the following field rates: 180 kg nitrogen·ha − 1 , 90 kg phosphorus·ha − 1 , and 240 kg potassium·ha − 1 , each pot received a total of 1.57 g urea, 1.92 g potassium sulfate, and 3.0 g calcium superphosphate during the growing season. The calcium superphosphate was applied as a basal fertilizer, while urea and potassium sulfate were applied in a ratio of 1:3:2 at the seedling, flowering, and early fruiting stages, respectively, through fertigation. Irrigation method The irrigation system consists of buried drip emitters, with an emitter working pressure of 0.10 Mpa. Standard drip irrigation treatments received water through the standard supply system, while for aerated treatments we utilized a Venturi air injector (Mazzei air injector 684, Mazzei Corp. USA) to provide oxygenation. Soil moisture content was monitored using a handheld Time Domain Reflectometry (TDR) device. When the soil moisture content dropped to approximately 60% of field capacity, irrigation was performed on the base of evaporation rate from a standard evaporation pan. The amount of irrigation water was calculated using the following Eq. ( 1 ) $$W=A \cdot {E_P} \cdot {K_P}$$ 1 Where W —Single irrigation amount (L); A —Area of the planting pot (m 2 ) (which was 0.038 m 2 in this study); E P —Evaporation from the pan between two irrigation events (mm); K P —Pan coefficient, taken as 0.8. 2.3 Sample collection and measurement After the tomatoes matured on January 21st, 2025, the plants were carefully removed from the soil. The soil samples were air-dried naturally and used to determine the soil physicochemical properties and phosphatase activity. The tomato plants were washed with deionized water and then separated into aboveground and underground parts. The fresh roots were scanned, weighed, and then a portion of about 1 cm of root segments was collected to determine the mycorrhizal colonization rate. After weighing the fresh root mass, the samples were blanched at 105°C for 30 minutes, followed by drying at 60°C to a constant weight. The dry weight was recorded, and the samples were then ground and passed through a 0.149 mm sieve. The phosphorus contents in both aboveground and underground parts of the plant were measured using the H 2 SO 4 -H 2 O 2 digestion method and molybdenum-antimony colorimetry. The phosphorus accumulation was calculated, and the phosphorus content in each part was determined using an AA3 Flow Analyzer. Rhizosphere soil samples were collected to measure available phosphorus and total phosphorus. 2.3.1. Mycorrhizal colonization rate : The root staining method was consists of improved acetic acid ink staining technique (Vierheilig et al. 1998). The colonization rate was calculated using the method widely adopted by the Arbuscular Mycorrhizal Fungi Germplasm Resource Bank (BGC), which determines the overall colonization rate as described by Biermann and Linderman (1981): $${\text{Mycorrhizal Colonization Rate}}(\% )=\frac{{\sum {(0 \times {S_i}+10 \times {S_i}+20 \times {S_i}+ \cdots 100 \times {S_i})} }}{{{\text{Total EquationNumber of root segments observed}}}}$$ 2 Under the microscope, the mycorrhizal colonization status of each root segment was examined (e.g., points of penetration, hyphae, arbuscules, vesicles, etc.). The colonization rate was scored according to the infection percentage, with values of 0, 10, 20, 30, …, 100, where 10 represents the first level of infection. For each segment, the number of root segments infected at each level (Si) was recorded. The mycorrhizal colonization rate of the sample was calculated by dividing the sum of product of the number of root segments and their respective levels by the total number of observed segments (Eq. 2 ). 2.3.2 Soil phosphorus fractionations : The phosphorus fractionations in the soil were determined using the modified Hedley phosphorus fractionation method (Tiessen and Moir, 1993). In brief, 0.5 g of air-dried soil passed through a 100-mesh sieve was weighed and treated with 30 mL deionized water (containing anion exchange resin) to extract water-soluble phosphorus (Resin-Pi) using the molybdenum-antimony colorimetric method. Subsequently, the soil was sequentially extracted with 0.5 M NaHCO 3 , 0.1 M NaOH, and 1.0 M HCl (D-HCl-Pi), with the C-HCl-P fraction extracted by heating in a water bath. Residual phosphorus (Residual-P) was extracted using H 2 SO 4 -H 2 O 2 digestion. Finally, the fractions were quantified using the molybdenum-antimony colorimetric method. 2.3.3 Soil phosphatase activity : Soil acid phosphatase activity was measured using the p-nitrophenyl phosphate disodium (PNPP) colorimetric method (Veloso et al. 2023; Tabatabai and Bremner, 1969). Accurately weighed 0.20 g of soil was passed through a 0.18 mm sieve and placed into a 10 mL round-bottom plastic centrifuge tube. Later on, 0.2 mL toluene was added, vortex to mix, and left to stand for 15 minutes. Then, 4 mL phosphate buffer and 1 mL of 0.05 M p-nitrophenyl phosphate disodium solution (prepared in phosphate buffer) was added. After capping, it was mixed thoroughly and incubated at 37°C for 1 hour. After incubation, shake the mixture, and added 1 mL of 0.5 M CaCl 2 and 8 mL of 0.5 M NaOH, vortex to mix, and centrifuge at 2500 rpm for 5 minutes. Next, we transferred 5 mL of the supernatant to a new tube and centrifuge at 3800 rpm for 5 minutes. Finally, we measured the absorbance at 410 nm. The acid phosphatase activity in the soil is expressed as the micrograms of p-nitrophenol per gram of soil per unit time, with units of mg·g − 1 ·d − 1 . 2.3.4 Root morphology parameters : After washing the roots with deionized water, the clean roots were placed in a transparent tray with a depth of 10 mm. Using forceps, the roots were gently spread out to ensure full extension. The roots were then scanned using a root scanner (Epson Expression 1600 Pro, Japan). The scanned root images were analyzed with the WinRHIZO image analysis system (Regent, Canada) to measure root morphology parameters such as total root length, root volume, and root surface area. 2.3.5 Crop phosphorus accumulation and phosphorus use efficiency : ( 1 ) Phosphorus accumulation in plants : The root, stem, leaf, and fruit samples were washed with clean water, then placed in an oven at 105°C for 15 minutes for blanching. Afterward, the samples were dried at 75°C to a constant weight and weighed. After weighing, the samples were ground through a 0.15 mm-mesh sieve and stored in bags for further analysis. The total phosphorus content in each organ was determined using the H 2 SO 4 -H 2 O 2 digestion method, and phosphorus accumulation in each organ was calculated using a flow analyzer (AA3, SEAL Analytical, Germany). ( 2 ) Mycorrhizal growth response (MGR) and mycorrhizal phosphorus uptake response (MPR) : The MGR and MPR were used to assess the impact of AMF symbiosis on tomato growth and phosphorus uptake in each treatment (Veiga et al. 2011). The MGR was calculated as follows: $$N{M_{mean}}AM,{\text{ }}MGR=\left( {\frac{{AM}}{{N{M_{mean}}}} - 1} \right) \times 100$$ 4 Where NM mean represents the average biomass of tomato plants in the non-AMF treatment group, and AM is the biomass of the tomato plants in each AMF treatment group. The same method applies for calculating MPR . ( 3 ) Phosphorus acquisition efficiency (PAE) and Phosphorus use efficiency (PUE) : PAE and PUE were used to characterize the phosphorus efficiency of the tested tomato plants (Deng et al. 2018; Han et al. 2022). The calculation formulas are as follows: $$PAE=\frac{{{P_{plant}}}}{{{W_{root}}}}$$ 5 $$PUE=\frac{Y}{{{P_{\text{a}}}}}$$ 6 Where, PAE represents the ability of the plant to acquire phosphorus from the soil, in mg·g − 1 ; P plant represents the total phosphorus uptake by the plant in mg·pot − 1 ; W root represents the dry weight of the roots in g·pot − 1 . While, PUE represents the ability of the plant to utilize the acquired phosphorus to generate biomass or yield in kg·kg − 1 ; Y represents the tomato yield in kg·ha − 1 ; P a represents the total phosphorus absorbed by plants per hectare in kg·ha − 1 . 2.4 Data analysis Prior to data analysis, outliers were removed, and the data were tested for normality and homogeneity of variance. A multivariate analysis of variance (Multivariate ANOVA) was conducted to examine the individual and interactive effects of irrigation water DO and AMF inoculation modes on tomato root morphology, yield, phosphorus accumulation, and soil phosphorus fractions. Duncan's post-hoc multiple comparisons were performed for significance testing, with a significance level set at 0.05. All data were analyzed using SPSS 25.0 (IBM, Armonk, NY, USA) statistical software. Structural equation modeling (SEM) was constructed in SmartPLS 4.1.1.4 (SmartPLS, Hamburg, Germany) to evaluate the direct and indirect effects of "fungal colonization rate, phosphatase activity, and root morphology" on soil phosphorus activation and tomato phosphorus utilization. Graphs were generated using Origin 2021 software (OriginLab Corporation, Northampton, MA, USA). 3. Results and analysis 3.1 Arbuscular mycorrhizal fungi colonization Aerated drip irrigation (ADI) significantly affected the mycorrhizal colonization rate of arbuscular mycorrhizal fungi (AMF) in greenhouse grown tomatoes ( p < 0.05) (Fig. 1 ). The results revealed that non-inoculated treatments (C and A) had 0% colonization rate, indicating that the soil sterilization was effective prior to the experiment and that the system was free from background AMF contamination. Under standard drip irrigation condition (C), the highest colonization rate (20.32%) was observed in the Rhizophagus intraradices (RI) inoculation treatment, which was 14.86% higher than the inoculation of Funneliformis mosseae (FM) treatment, while the mixed inoculation (CFMRI) treatment had the lowest (14.68%) colonization rate. Under aerated conditions, the ARI treatment had the highest colonization rate of 26.95%, which was 32.66% higher than the CRI treatment and represented the highest rate across all treatments. The next highest inoculation rate (20.60%) was observed in AFM treatment followed by the AFMRI treatment (19.44%). Overall, the RI inoculation treatment exhibited a higher colonization rate under both irrigation conditions, while no significant difference was found between the RI and RIFM mixed inoculation treatments. This suggests that the RI inoculant was more effective at colonizing tomatoes in this system compared to FM, which may be attributed to differences in host adaptation between the strains. Additionally, the colonization rate of mixed inoculation treatment is lower than that of RI inoculation treatment alone, indicating that there may be competition or inhibition effect between mixed inoculation agents. Furthermore, the mycorrhizal colonization rate under ADI conditions was generally higher than that of standard drip irrigation, which may be related to the improved soil aeration by ADI, facilitating the germination and infection of fungal spores. The ADI system regulation of the rhizosphere oxygen environment, enhancing AMF activity, may further regulate phosphorus transformation and utilization, as well as tomato yield formation. This combined use of ADI and AMF and their effect is defined as the "oxygen-mycorrhiza synergy". 3.2 Root morphology characteristics The results of root morphology parameters under different treatments are given in Fig. 2 . It was observed that, ADI significantly upregulated the total root length, root surface area, and root tip number in tomatoes ( p < 0.05). The overall trends of total root length, root surface area, root volume, and root tip number were consistent, with the ARI treatment showing the highest values, the C treatment without inoculation showing the lowest. While, A treatment without inoculation and the CRI treatment with single inoculation were falling between these two high and low extremes (ARI > A ≈ CRI > C). No significant patterns were observed in the average root diameter, and neither ADI nor AMF treatments had a significant impact on this parameter. Under standard drip irrigation conditions (C), inoculation with Rhizophagus intraradices (CRI) significantly improved root morphology: total root length (980.02 cm), root surface area (383.82 cm 2 ), root volume (11.97 cm 3 ), and root tip number (2194) which were increased by 13.24%, 17.51%, 21.83%, and 13.25%, respectively, as compared to their control treatment (C). Inoculation with Funneliformis mosseae (CFM) also showed some enhancement in these four root parameters ( p > 0.05), but the increases were less than those observed with the RI inoculation. The mixed inoculation (CFMRI) treatment did not demonstrate any synergistic advantage; its root length, root surface area, and root tip number were similar to those of the C treatment without inoculation. However, in terms of average root diameter, the mixed inoculation treatment showed the highest value of 1.27 mm, which was a 5.85% increase compared to the control (C). ADI enhances the positive effects of AMF inoculation. Except for the average root diameter, the root morphology parameters under the RI inoculation treatment (ARI) in aerated conditions were the highest across all treatments: total root length (1211.82 cm) was 23.65% higher than the standard drip irrigation CRI treatment; root surface area (460.42 cm 2 ) increased by 19.96%; root volume (14.11 cm 3 ) increased by 17.90%; and the number of root tips (2714) increased by 23.70% compared to CRI. Under aerated conditions, inoculation with Funneliformis mosseae (FM) and the mixed inoculation (FMRI) also showed improvements in root length, surface area, and root tip number compared to standard drip irrigation, but the increases were less significant than those observed with RI inoculation. Furthermore, the mixed inoculation treatment did not perform well under both irrigation modes except for the average root diameter being similar to or even slightly lower than those under FM treatment. 3.3 Soil phosphorus fractionations and soil phosphatase activity The modified Hedley phosphorus fractionation method (Tiessen and Moir 1993) divides soil phosphorus into 9 fractions. In this study, we classified soil phosphorus into three categories based on its ease of absorption by plants: ( 1 ) Easily available phosphorus, which includes resin-exchangeable phosphorus (Resin-P), and two NaHCO 3 -extracted phosphorus fractions (NaHCO 3 -Pi, NaHCO 3 -Po); ( 2 ) Moderately available phosphorus, which includes two NaOH-extracted phosphorus fractions (NaOH-Pi, NaOH-Po); ( 3 ) Insoluble phosphorus, which includes phosphorus extracted with diluted hydrochloric acid (D-HCl-P), two phosphorus fractions extracted with concentrated hydrochloric acid (C-HCl-Pi, C- HCl-Po), and residual phosphorus (Residual-P). The changes in each phosphorus fraction under different treatments are summarized as follows. 3.3.1 Easily available phosphorus (Easily available P) The results for directly and easily available phosphorous for plant uptake or rapidly mineralized into available phosphorus including Resin-exchangeable phosphorus (Resin-P), NaHCO 3 -extracted inorganic phosphorus (NaHCO 3 -Pi), and some NaHCO 3 -extracted organic phosphorus (NaHCO 3 -Po) are given in in Fig. 3 . We found that aeration without inoculation significantly increased (42.03%) the Resin-P content in soil ( p < 0.01), showing a value of 18.99 mg·kg − 1 as compared to the control group (C) without inoculation having a value of 13.37 mg·kg − 1 . The CRI, CFM, and mixed inoculation (CFMRI) treatments increased Resin-P by 53.52%, 51.66%, and 25.83%, respectively than that of irrigation without aeration and inoculation (C). On the other hand, ARI, AFM, and AFMRI treatments increased Resin-P by 42.78%, 26.63%, and 22.14%, respectively, as opposed to A treatment without inoculation. Inoculation with RI, FM, and the RI + FM mixed inoculation, combined with ADI, further increased Resin-P by 32.09%, 18.59%, and 37.87%, respectively, compared to the only inoculation treatments (CRI, CFM and CFMRI). In summary, applying arbuscular mycorrhizal inoculation under aerated drip irrigation significantly increased soil Resin-P, and their combined use demonstrated a positive synergistic effect, specifically achieved under combination of ADI and the RI inoculant (ARI). NaHCO 3 -Pi is an active inorganic phosphorus extracted by sodium bicarbonate solution, which is mainly in an easily adsorbed form (phosphorus weakly bound to aluminum, iron, and calcium). Compared to the NaHCO 3 -Pi content (37.21 mg·kg − 1 ) in the C treatment (without inoculation and aeration treatment), both aerated drip irrigation and inoculation with fungi significantly affected the NaHCO 3 -Pi content in the soil ( p < 0.01). The value for NaHCO 3 -Pi content in aerated treatment without any inoculation (A) was 44.67 mg·kg − 1 , which was 20.04% high compared to control group (C). The CRI, CFM, and mixed inoculation (CFMRI) treatments increased NaHCO 3 -Pi by 82.41%, 57.98%, and 42.39%, respectively, compared to the control group (C). While under aerated irrigation, ARI, AFM, and AFMRI treatments increased NaHCO 3 -Pi by 76.07%, 49.13%, and 43.95%, respectively, compared to aerated treatment (A). When inoculation was done with RI, FM, or the RI + FM under ADI, NaHCO 3 -Pi was further increased by 15.87%, 13.32%, and 21.36%, respectively, compared to CRI, CFM, and CFMRI treatments. Overall, AMF inoculation under aerated drip irrigation increased the soil NaHCO 3 -Pi content, and their combined use demonstrates a significant synergistic effect, with the maximum increase observed in the ADI + RI inoculant combination (ARI). The NaHCO 3 -Po is mainly an easily mineralizable organic phosphorus and is an important source of available phosphorus in the soil. In contract to NaHCO 3 -Pi, the value for NaHCO 3 -Po (45.33 mg·kg − 1 ) in the aerated treatment (A) was 11.01% less as compared to control group value (50.94 mg·kg − 1 ). The CRI, CFM, and mixed inoculation (CFMRI) treatments reduced NaHCO 3 -Po by 18.71%, 16.65%, and 7.5%, respectively, compared to the control group (C). The ARI, AFM, and AFMRI treatments reduced NaHCO 3 -Po by 30.25%, 17.36%, and 13.99%, respectively, compared to aerated treatment without inoculation (A). When combined RI, FM, or the RI + FM mixed inoculation with ADI, NaHCO 3 -Po was further reduced by 23.64%, 11.78%, and 17.26%, respectively, compared to inoculation treatments without aeration. Generally, AMF inoculation under aerated drip irrigation led to a reduction in NaHCO 3 -Po, with the maximum reduction observed in the ADI + RI inoculant combination (ARI). Overall, both ADI and AMF technologies increased the total easily available phosphorus fractions (Resin-P, NaHCO 3 -Pi, and NaHCO 3 -Po) in the soil. Compared to the easily available phosphorus content in the control soil (101.52 mg·kg − 1 ), aerated drip irrigation (A) had 35.3% increase in this value. Inoculation with AMF also showed similar effects, with RI, FM, and mixed inoculation increasing the total easily available phosphorus by 27.86%, 19.70%, and 15.18%, respectively as compared to CRI, CFM, and CFMRI treatments. Under oxygen-mycorrhiza synergy conditions, this effect was more pronounced, such as ARI, AFM, and AFMRI treatments increased total easily available phosphorus by 35.32%, 26.20%, and 24.59%, respectively, compared to the C treatment. The ADI + RI inoculant (ARI) combination achieved the maximum increase in efficiency. 3.3.2 Moderately available phosphorus (Moderately available P) The NaOH-extracted inorganic phosphorus (NaOH-Pi) and organic phosphorus (NaOH-Po) were categorized as moderately available phosphorus, which lies between easily available phosphorus and insoluble phosphorus. These fractions can be released over a longer time scale through desorption and mineralization. The results obtained from application of irrigation modes and AMF inoculation in tomato plants are presented in Fig. 4 . We found that there were no statistical differences among various treatments for NaOH-Pi contents. Generally, the NaOH-Pi value in control group (C treatment) was 84.03 mg·kg − 1 , while it was 81.09 mg·kg − 1 in aerated drip irrigation treatment (A), a 3.5% decrease compared to control group (C). The CRI, CFM, and mixed inoculation (CFMRI) treatments reduced NaOH-Pi by 9.13%, 5.95%, and 7.14%, respectively, compared to the control group (C). Further, ARI, AFM, and AFMRI treatments reduced NaOH-Pi by 8.63%, 6.13%, and 2.5%, respectively, that of alone aerated treatment (A). Besides, inoculation with RI, FM in aerated drip irrigation further reduced the NaOH-Pi by 2.98% and 3.69% as opposed to CRI and CFM treatments respectively. NaOH-Po is primarily organic phosphorus bound with humus. In contrast to NaOH-Pi, the application of inoculation significantly affected the NaOH-Po contents under different irrigation modes. The NaOH-Po value was 61.16 mg·kg − 1 in the control group (C treatment), while this value was 56.48 mg·kg − 1 in alone aerated treatment (A), showing a 7.66% decrease compared to the control group (C). The CRI, CFM, and mixed inoculation (CFMRI) treatments reduced NaOH-Po by 22.15%, 20.74%, and 10.66%, respectively than control group (C). Considering the aerated irrigation treatments, ARI, AFM, and AFMRI treatments reduced NaOH-Po by 33.04%, 29.2%, and 15.79%, respectively, compared to alone aerated treatment (A). Combining RI, FM, or the RI + FM mixed inoculation with aerated drip irrigation further reduced NaOH-Po by 20.57%, 17.52%, and 12.96%, respectively, compared to the CRI, CFM, and CFMRI treatments. The combined use of ADI and AMF showed a significant synergistic effect, with the best results observed in the ADI + RI inoculant combination (ARI). Overall, both ADI and AMF technologies reduced the total moderately available phosphorus contents in the soil. Compared to the control group values (145.19 mg·kg − 1 ), soil moderately available P content (137.95 mg·kg − 1 ) in aerated drip irrigation (A) was reduced by 5.25%. Inoculation with AMF also showed similar effects: for example, under the standard drip irrigation condition, RI, FM, and mixed inoculation reduced the moderately available P content by 14.61%, 12.18%, and 8.62%, respectively, as compared to the C treatment. Under oxygen-mycorrhiza synergy conditions, this effect was more pronounced, with ARI, AFM, and AFMRI treatments reduced the total contents of moderately available phosphorous by 22.93%, 20.04%, and 12.79%, respectively, compared to the C treatment. 3.3.3 Insoluble phosphorus (Insoluble P) Insoluble forms of phosphorus (D-HCl-P, C-HCl-Pi/Po, and Residual-P) are typically not directly available to plants and can only be released as available phosphorus through long-term soil changes or microbial activity (Xu et al. 2020). The results regarding effects of different irrigation modes and mycorrhizal fungi inoculation on insoluble P fractions are given in Fig. 5 . D-HCl-P is a phosphorus fraction extracted with diluted hydrochloric acid, primarily in the form of apatite and calcium-bound phosphorus. We found that, the D-HCl-P value was 252.57 mg·kg − 1 in control group (C treatment), while aerated drip irrigation treatment (A) showed a value of 237.45 mg·kg − 1 , a 5.99% decrease compared to the control group (C). The CRI, CFM, and mixed inoculation (CFMRI) treatments reduced D-HCl-P by 11.96%, 12.83%, and 8.36%, respectively, compared to control group (C). When inoculation was done with RI, FM, or the RI + FM mixed inoculation under aerated drip irrigation, it was found that D-HCl-P was further reduced by 10.87%, 6.74%, and 6.03% than CRI, CFM, and CFMRI, respectively. In summary, both aerated drip irrigation and AMF inoculation led to a reduction in soil D-HCl-P ( p > 0.05), with no significant synergistic effect when combined. C-HCl-Pi is stable inorganic phosphorus extracted with concentrated hydrochloric acid. There was no significant effect of aeration and fungi inoculation on C-HCl-Pi. Overall, it was observed that inoculation increased the C-HCl-Pi under both aerated and non-aerated condition. However, aerated drip irrigation has higher values for C-HCl-Pi than non-aerated irrigation mode ( p > 0.05). C-HCl-Po is stable organic phosphorus extracted with concentrated hydrochloric acid. We noticed that the aerated drip irrigation treatment (A) showed a 14.48% decrease in C-HCl-Po value compared to the control group (C). Inoculation with fungi also reduced the C-HCl-Po, such as CRI, CFM, and mixed inoculation (CFMRI) treatments have 19.15%, 13.55%, and 8.15% less C-HCl-Po compared to the control group (C) respectively. Besides, ARI, AFM, and AFMRI further reduced C-HCl-Po by 25.37%, 19.54%, and 13.64%, respectively as opposed to CRI, CFM, and CFMRI respectively. Briefly, synergistic effect of both aerated drip irrigation and AMF inoculation reduced the soil C-HCl-Po content ( p < 0.05), with the highest reduction observed in ADI + RI inoculant combination (ARI). Residual-P is soil residual phosphorus extracted by strong acid digestion. The Residual-P value in the control group (C treatment) was 104.49 mg·kg − 1 and was 101.28 mg·kg − 1 in aerated drip irrigation treatment (A), showing that aerated irrigation reduced Residual-P. The inoculation also notably minimized the Residual-P in soil, for example CRI, CFM, and mixed inoculation (CFMRI) treatments reduced Residual-P by 6.19%, 6.41%, and 10.78% respectively than control group (C). Furthermore, synergistic effect of ADI and AMF further decreased the Residual-P by 6.48%, 2.44%, and 3.52% than CRI, CFM, and CFMRI respectively ( p < 0.05). Overall, both ADI and AMF technologies slightly reduced the content of total insoluble phosphorus fractions in the soil. Compared to the control group contents (493.45 mg·kg − 1 ), soil insoluble P content in aerated drip irrigation (A) reduced by 3.55%. Inoculation with AMF showed similar decreasing effects, such as RI, FM, and mixed inoculation reduced the total insoluble P content by 5.18%, 6.74%, and 4.68%, respectively. Under oxygen-mycorrhiza synergy conditions, this effect in lowering the total insoluble phosphorus fraction was enhanced, with ARI, AFM, and AFMRI treatments reducing it by 11.94%, 9.63%, and 6.69% respectively compared to the control group. Further, ADI + RI inoculant (ARI) combination showed the best effect. 3.3.4 Soil acid phosphatase activity The results regarding soil acid phosphatase (Acid Phosphatase, ACP) activity measured in µmol p-nitrophenol·g − 1 dry soil·h − 1 are given in Fig. 6 . The findings revealed that ACP value in the control group (C treatment) was 5.12 µmol·g − 1 ·h − 1 . While in aerated drip irrigation treatment (A), the ACP value was 7.2 µmol·g − 1 ·h − 1 , which represented a 40.63% increase compared to the control group (C). The CRI, CFM, and mixed inoculation (CFMRI) treatments increased ACP activity by 54.81%, 25.98%, and 12.96%, respectively, compared to C treatment. Furthermore, the corresponding values for ACP activity under ARI, AFM, and AFMRI were high by 29.38%, 26.05%, and 35.22%, respectively, than CRI, CFM, and CFMRI inoculation treatments. Overall, combined use of ADI and AMF inoculation exhibited a synergistic effect, with the maximum increase observed in the ADI + RI inoculant combination (ARI). 3.4 Phosphorus accumulation in different plant organs, yield, and phosphorus use efficiency The effects of ADI and AMF inoculation on phosphorus accumulation in the different organs of tomato plants are given in Fig. 7 . Both ADI and AMF inoculation significantly affected phosphorus accumulation in the root, stem, leaf, and tomato fruits ( p fruit > stem > root. The total phosphorus accumulation in aerated drip irrigation treatment (A) was 216.44 mg.pot − 1 , representing a 30.82% increase as compared to control group (C) value (165.45 mg·pot − 1 ). The CRI, CFM, and mixed inoculation (CFMRI) treatments increased total phosphorus accumulation by 22.94%, 15.09%, and 9.47%, respectively than the treatment without any inoculation and aeration (C). Besides, the corresponding values for total phosphorus accumulation under ARI, AFM, and AFMRI were further increased under aerated irrigation by 34.38%, 25.73%, and 21.37%, respectively as compared to CRI, CFM, and CFMRI. In summary, both aerated drip irrigation and AMF inoculation showed a significant synergistic effect which increased the total phosphorus accumulation in tomato plants, with the highest total phosphorus accumulation observed in the ADI + RI inoculant combination (ARI). Under the ARI treatment, the phosphorus accumulation in the stem, leaf, and fruit of tomatoes increased by 55.42%, 72.39%, and 80.66%, respectively, as opposed to the control group (C). The effects of ADI and AMF inoculation on tomato yield (fresh weight) and mycorrhizal attributes are presented in Table 3 . We noticed that ADI significantly improved tomato yield ( p AFM > AFMRI > A > CRI > CFM > CFMRI > C. The aerated drip irrigation treatment (A) yielded 57.08 t·ha − 1 , which was 26.53% more yield as compared to the control treatment (C) that produced 45.12 t·ha − 1 . Inoculation without aerated irrigation improved yield by 16.58%, 14.36%, and 10.83% in CRI, CFM, and CFMRI respectively than control (C). While ARI, AFM, and AFMRI treatments increased yield by 26%, 14.24%, and 10.76%, respectively, compared to the aerated treatment without inoculation (A). Overall, both aerated drip irrigation and AMF inoculation promoted an increase in tomato yield, with the highest yield observed in the ADI + RI inoculant combination (ARI). Table 3 Tomato yield, mycorrhizal effects, and phosphorus use efficiency. Treatment Yield (t·ha − 1 ) MGR (%) MPR (%) PUE (kg·kg − 1 ) PAE (mg·g − 1 ) C 45.12 ± 3.87d - - 1038.06 ± 26.16a 28.91 ± 2.86c CRI 52.6 ± 2.08cd 15.19 ± 3.02b 18.52 ± 3.41cd 985.16 ± 73.04a 34.9 ± 5.28bc CFM 51.6 ± 2.3cd 14.05 ± 3.74b 12.92 ± 4.12cd 1030.46 ± 40.89a 29.89 ± 4.49c CFMRI 50 ± 2.28cd 9.91 ± 4.46b 7.93 ± 4.03d 1053.81 ± 61.07a 25.88 ± 2.69c A 57.08 ± 5.42bc - - 1004.19 ± 68.71a 35.01 ± 3.1bc ARI 71.93 ± 7.7a 35.16 ± 7.29a 43.31 ± 2.53a 997.27 ± 59.38a 47.91 ± 7.93a AFM 65.21 ± 0.47ab 30.28 ± 0.63a 30.83 ± 2.19b 1035.82 ± 29.25a 45.37 ± 7.93ab AFMRI 63.22 ± 3.31ab 26.69 ± 4.16a 24.12 ± 6.75bc 1096.98 ± 51.28a 37.19 ± 5.31abc Data are presented as mean ± standard deviation. Values with the same lower case were not significant at p < 0.05 among different treatments. C represents standard drip irrigation, and A represents aerated drip irrigation. RI refers to the treatment with Rhizophagus intraradices (RI) inoculation only; FM refers to the treatment with Funneliformis mosseae (FM) inoculation only; FMRI refers to the mixed inoculation of FM and RI. Additionally, aeration significantly enhanced the mycorrhizal growth response ( MGR ) and mycorrhizal phosphorus uptake response ( MPR ) in studied tomatoes ( p < 0.01). The analysis showed that under standard drip irrigation conditions (C), the CRI treatment had the highest MGR and MPR at 15.19% and 18.52%, respectively, which were higher than the 14.05% and 12.92% in the CFM treatment, while the mixed inoculation (CFMRI) treatment had the lowest MGR and MPR at 9.91% and 7.93%, respectively. Same trend was found under ADI conditions, with the ARI treatment showing the highest MGR and MPR at 35.16% and 43.31%, respectively. Hence, RI inoculation showed high mycorrhizal effects under both irrigation conditions, indicating that RI inoculation contributed more to tomato dry matter accumulation and phosphorus uptake compared to FM, while aeration significantly enhanced this inoculation effect. The phosphorus use efficiency indicators ( PAE and PUE ) for tomatoes are given in Table 3 . The aerated drip irrigation treatment (A) had PAE value of 35.01 mg·g − 1 , showing 21.12% increase as compared to control treatment (C) which has PAE value of 28.91 mg·g − 1 . The CRI and CFM treatments increased PAE by 20.75% and 3.39%, respectively, than control (C). The ARI, AFM, and AFMRI treatments increased PAE by 36.86%, 29.59%, and 6.24%, respectively as opposed to their control treatment without any inoculation (A). Briefly, both aerated drip irrigation and AMF inoculation significantly increased PAE , and their combined use demonstrated a significant synergistic effect ( p 0.05). The aerated drip irrigation treatment (A) had 3.26% decrease in PUE than control treatment (C). The CRI, CFM, and mixed inoculation (CFMRI) treatments affected PUE in tomato by -5.1%, -0.73%, and 1.52%, respectively, compared to C treatment. The RI, FM, and RI + FM inoculation treatments under aerated drip irrigation increased PUE by 1.23%, 0.52%, and 4.1%, respectively, compared to the CRI, CFM, and CRIFM. Overall, aerated drip irrigation slightly improved PUE , while the effect of AMF inoculation on PUE was inconsistent ( p > 0.05), because the best performance for PUE was observed in the ADI + mixed inoculant combination (AFMRI) instead of ARI. 3.5 Structural equation model of phosphorus accumulation and yield formation in greenhouse tomato A structural equation model (SEM) was constructed to analyze the synergistic effects of aerated drip irrigation (ADI) and arbuscular mycorrhizal fungi (AMF) inoculation on soil phosphorus transformation, root morphology development, plant phosphorus accumulation, and tomato yield regulation (Fig. 8 ). The variance explanation (R 2 ) for each endogenous variable was as follows: soil phosphatase activity (61.0%), soil inorganic phosphorus (44.4%), soil organic phosphorus (46.1%), root morphology development (44.9%), plant phosphorus accumulation (68.4%), and yield (86.1%). These results indicate that the model has good explanatory power and integrates the key processes affecting tomato yield formation. The path analysis of the structural equation model revealed the following: ( 1 ) Drivers of soil phosphatase activity : Dissolved oxygen in irrigation water (path coefficient = 1.219, p < 0.001) and AM fungi (path coefficient = 0.517, p < 0.001) both significantly and positively influenced soil phosphatase activity, indicating that both factors synergistically promoted the activation of phosphatase. ( 2 ) Regulation of phosphorus fractions by phosphatase : We found that soil phosphatase activity significantly and positively drove soil inorganic phosphorus (path coefficient = 0.666, p < 0.001), while negatively regulated soil organic phosphorus (path coefficient = -0.679, p < 0.001). This reflects that phosphatase accelerates organic phosphorus mineralization and enriches inorganic phosphorus. ( 3 ) Response of root morphology development : Dissolved oxygen in irrigation water (path coefficient = 1.030, p < 0.001) and AM fungi (path coefficient = 0.452, p < 0.01) inoculation both significantly and positively shaped root morphology development, indicating that ADI may improve root morphology development by alleviating root hypoxic stress, while AM fungi might optimize root morphology through hyphal extension and signaling regulation. ( 4 ) Factors influencing plant phosphorus accumulation : Soil organic phosphorus contents were negatively associated with plant phosphorus accumulation (path coefficient = -0.451, p < 0.05), while root morphology development significantly promoted phosphorus accumulation (path coefficient = 0.365, p 0.05). These three factors together explained 68.4% (R 2 = 0.684) of the variation in plant phosphorus accumulation. Briefly, oxygen-mycorrhiza synergy might promote phosphorus accumulation by enhancing soil phosphatase activity, accelerating organic phosphorus mineralization, and regulating inorganic phosphorus supply, as well as optimizing root morphology to directly promote phosphorus absorption. ( 5 ) Dominant pathways in yield formation : Plant phosphorus accumulation significantly and positively determined yield (path coefficient = 0.911, p 0.05), indicating that yield improvement mainly relies on the indirect effects of efficient phosphorus accumulation. The model reliability and validity tests showed that most of the latent variables had Cronbach's α > 0.8, while all variables have average variance extracted (AVE) > 0.5, and composite reliability (CR) > 0.7, confirming that measurement model has good internal consistency and convergent validity. In conclusion, ADI and AMF synergistically improve soil phosphatase activity, promote organic phosphorus mineralization and inorganic phosphorus supply, which leads to optimize root morphology, ultimately increasing plant phosphorus accumulation and enhancing yield. Further, soil phosphatase plays a central role in phosphorus transformation, and plant phosphorus accumulation is a key mediator of yield formation. 4. Discussion This study systematically explored the synergistic effects of aerated drip irrigation (ADI) and AMF inoculation on soil phosphorus transformation, root morphology, phosphorus accumulation, and yield. The results showed that the combination of ADI and Rhizophagus intraradices (RI) inoculation (ARI treatment) exhibited significant synergistic effects in enhancing AMF colonization, improving root morphology, promoting soil phosphorus transformation, and increasing phosphorus uptake and yield in tomatoes. This study therefore introduces the concept of "Oxygen-Mycorrhiza Synergy". 4.1 Oxygen-mycorrhiza synergy on soil phosphorus fractions and availability The availability of soil phosphorus directly impacts plant phosphorus uptake and growth. The transformation of phosphorus fractions is a key process in enhancing phosphorus availability. Based on the Hedley phosphorus fractionation system, this study found that the effects of ADI and AMF inoculation on soil phosphorus fractions followed the gradient pattern of "easily available phosphorus > moderately available phosphorus > insoluble phosphorus". The AMF inoculation increased easily available phosphorus contents in soil (Resin-P and NaHCO 3 -Pi), while decreasing the organic phosphorus components (such as NaHCO 3 -Po and NaOH-Po). However, the short-term impact of AMF on stable phosphorus fractions in the soil (HCl-P, Residue-P) was minimal. Phosphatases play a critical role in the microbial-mediated conversion of insoluble phosphorus (Lidbury et al. 2022), and this transformation indicates that AMF promoted organic phosphorus mineralization through the secretion of acid phosphatase, thereby enhancing phosphorus availability in the soil. This phenomenon aligns with the results observed in our study, where organic phosphorus decreased, and available phosphorus increased, as AMF hyphae exhibit higher acid phosphatase activity in the presence of organic phosphorus (Huang et al. 2023). The oxygen-mycorrhiza synergy amplified this transformation effect and significantly altered the distribution of soil phosphorus. Under the ARI treatment, soil acid phosphatase activity was 29.38% higher than that in the CRI treatment, while easily available phosphorus increased by 5.83%. Besides, moderately available phosphorus and insoluble phosphorus decreased by 9.74% and 7.13% respectively. The possible reasons for these changes could be: ( 1 ) ADI may have selectively enriched microbial groups related to phosphorus transformation (such as Proteobacteria and Acidobacteria) by altering soil redox conditions, further optimizing phosphorus cycling pathways (Gross et al. 2020; Kim et al. 2021); ( 2 ) Hypoxia could reduce the abundance of phosphate-dissolving bacteria such as Pseudomonas and Bacillus (Cui et al. 2022), and these bacteria usually dissolve insoluble phosphate by secreting organic acids and enzymes (Ouyang et al. 2021). As ADI increased dissolved oxygen in the irrigation water and soil porosity, thereby improving soil gas exchange, which likely enhanced the phosphorus-solubilizing bacteria and AMF metabolic activity. The organic acids (such as citric acid and oxalic acid) secreted by phosphorus-solubilizing bacteria lower soil pH, chelate metal ions like Fe 3+ and Al 3+ , and dissolve certain phosphates (e.g., Fe-P, Al-P) (Pang et al. 2024; Aliyat et al. 2022). Meanwhile, AMF upregulates the expression of genes related to host plant acid phosphatase secretion, which increases soil acid phosphatase activity and accelerates organic phosphorus mineralization (Nopphakat et al. 2022; Pang et al. 2024). As a result, oxygen-mycorrhiza synergy enhanced the availability of phosphorus to tomatoes. This finding has significant implications for optimizing soil phosphorus management and improving phosphorus fertilizer use efficiency. 4.2 Effects of oxygen-mycorrhiza synergy on tomato-mycorrhizal symbiosis and phosphorus uptake Mycorrhizal symbiosis is an important pathway for plants to acquire mineral nutrients, especially under low phosphorus availability in soil. The colonization rate is a key indicator of the effectiveness of mycorrhizal symbiosis. In our study, ADI significantly increased tomato mycorrhizal colonization, particularly after inoculation with Rhizophagus intraradices (RI), which showed a 32.66% increase in colonization rate compared to conventional drip irrigation without aeration. This may be closely related to the improvement of rhizosphere oxygen conditions by ADI, which facilitated spore germination and hyphal invasion. Suitable soil oxygen concentrations are crucial for AMF growth and activity, as AMF are aerobic microorganisms whose metabolic activities depend on oxygen supply (Zhang et al. 2014). Xu et al. (2021) found that aeration significantly increased dissolved oxygen concentrations and AMF colonization in wetland ecosystems, promoting plant growth. In current study, intermittent aeration (4 hours·day − 1 ) might stabilized the aerobic environment, facilitating AMF colonization of roots. Furthermore, ADI improved soil aeration, promoted root respiration, and increased the secretion of carbohydrates (such as sucrose and glucose) from roots (Zheng et al. 2024), providing more carbon sources for AMF, thereby accelerating the formation of hyphal networks. Moreover, aeration may enhance the plant's ability to recognize and colonize AMF by modulating plant hormone balance (such as auxins and abscisic acid) (Chareesri et al. 2020). Notably, RI inoculation showed higher colonization efficiency than Funneliformis mosseae (FM) and the mixed inoculation (FM + RI), suggesting potential competition effects between different AMF strains (Berruti et al. 2016). Oxygen-mycorrhiza synergy also optimized tomato root morphology. The AMF inoculation remarkably improved plant nutrient uptake by changing root branching patterns and increasing the proportion of fine roots (Chandrasekaran 2022). In this study, soil aeration significantly influenced tomato root length, root surface area, and root tip number ( p < 0.05). Specifically, under the ARI treatment, the total root length, root surface area, and root branching increased by 23.65%, 19.96%, and 23.67%, respectively, compared to the CRI treatment. In contrast, the average root diameter ranged from 1.10 to 1.27 mm, with no significant differences observed between treatments. The effect of aeration on root diameter was not significant ( p > 0.05), which may be attributed to the fact that although ADI increased root volume, it also promoted root growth and elongation (as indicated by increased root tip number and total root length), thus exerting no significant impact on root diameter. However, it also suggests that tomato roots have a larger specific surface area, more branching, and finer root hairs, which facilitate better interaction with soil nutrients (Nascimento et al. 2021). Furthermore, ADI improved root development by enhancing soil oxygen supply, as oxygen is essential for root growth and metabolic activities (Roosta 2024; Wang et al. 2023b). The synergistic effects of ADI and AMF to amplify the improvement in root morphology might be attributed to ADI alleviating soil compaction and hypoxic stress, enhancing root vitality and respiration efficiency. The improved root structure provided more colonization sites for AMF, and the enhanced nutrient absorption capacity of AMF promoted denser lateral root development, thereby increasing surface area and forming a more efficient material exchange interface. The core mechanism of oxygen-mycorrhiza synergy lies in optimizing the rhizosphere microenvironment, activating fungal functional expression, and reshaping root growth patterns, ultimately affecting root absorption and supporting functions (Zeng et al. 2025; Wang et al. 2023a). Both aerated drip irrigation and AMF inoculation significantly increased tomato phosphorus accumulation, consistent with previous studies (Bowles et al. 2016). Under oxygen-mycorrhiza synergy, higher phosphorus availability and improved root morphology led to significant increases in phosphorus accumulation in tomato stems, leaves, and fruits. In this study, the ARI treatment showed a 34.38% higher phosphorus accumulation in fruit compared to CRI. Furthermore, the ARI treatment achieved the highest yield (71.93 t·ha − 1 ), a 36.75% increase compared to CRI. Similar findings were also reported by Leventis et al. (2021) in controlled experiments with tomato inoculation (Leventis et al. 2021). The reasons behind the increase in tomato productivity under combined use of ADI and AMF could be: AMF may break through the phosphorus-depleted zones in the rhizosphere with its hyphal networks, transferring phosphorus to the host plant and expanding the root nutrient absorption range (Wen et al. 2020); while aerated drip irrigation improves the soil microenvironment, enhancing AMF activity and colonization, thus interactively promoting phosphorus uptake by plants. It is important to note that different AMF inoculation modes contributed differently to tomato phosphorus accumulation and yield. The mycorrhizal effect of RI inoculation was significantly high than that of FM inoculant and mixed inoculation, which may be due to the stronger adaptability of the RI strain to the tested soil and oxygen environment. In mixed inoculation, the growth-promoting effect of FM and RI may be weakened or even antagonistic due to resource competition, niche overlap, or metabolic interference (Zhang et al. 2024), although still higher than the control group without inoculation. Additionally, symbiosis is often fine-tuned based on plant needs and surrounding conditions, typically through plant hormone signaling. Lidoy et al. (2023) compared the colonization ability of Funneliformis mosseae and Rhizophagus irregularis under hormone and salt stress conditions, concluding that colonization levels depend on the fungal genotype and stress conditions. In this study, soil aeration may have influenced the phosphorus signaling pathways of tomato plants by modulating soil oxygen stress, preferentially activating gene expression for symbiosis with RI, thereby enhancing the functional advantage of the specific fungal strain. This was reflected in the best symbiotic effect observed with the RI inoculant in tomato plants. 4.3 Mechanism involved in oxygen-mycorrhiza synergy to increase tomato yield and phosphorus acquisition The SEM revealed the potential mechanisms by which ADI and AMF synergistically enhance yield formation. The model showed that soil phosphatase activity is a key factor in phosphorus transformation and significantly positively influenced the soil inorganic phosphorus contents (path coefficient = 0.666). This finding is consistent with research suggesting that AMF increase phosphorus availability by modulating soil enzyme activity (Sheteiwy et al. 2021), and we notably found that oxygen-mycorrhiza synergy mainly promotes the conversion of organic phosphorus to inorganic phosphorus. In this study, oxygen-mycorrhiza synergy optimized phosphorus supply by enhancing phosphatase activity and improving the structure of soil phosphorus fractions. Simultaneously, it promoted tomato root development (path coefficient = 1.030) and enhanced the plant phosphorus uptake capacity (path coefficient = 0.365). These results indicate that the combined application of aerated drip irrigation and AMF not only promotes effective phosphorus absorption by enhancing soil phosphatase activity and inorganic phosphorus supply, but also further improved phosphorus accumulation and yield in tomatoes by optimizing root morphology. Moreover, the model also revealed the significant positive impact of phosphorus accumulation on tomato yield (path coefficient = 0.911), emphasizing the crucial role of phosphorus uptake in yield formation. Finally, our findings underscore the importance of integrating soil biological processes and plant physiological responses in optimizing nutrient management and crop production. 5. Conclusion This study emphasized the critical role of oxygen-mycorrhiza synergy in regulating availability of soil phosphorous via complex interactions involving mineralization, colonization, and enzymatic activities. We found that aeration can optimize the symbiotic relationship between tomato roots and AMF, thereby significantly increasing mycorrhizal colonization rate. The oxygen-mycorrhiza synergy notably improved acid phosphatase activity, promoted the conversion of soil active organic phosphorus to active inorganic phosphorus, and facilitates the transformation of insoluble phosphorus to available phosphorus. Simultaneously, both ADI and AMF optimized tomato root morphology, jointly promoting phosphorus accumulation in plants and ultimately increasing yield. In summary, the combined use of ADI and AMF has a significant synergistic effect in promoting phosphorus uptake in tomatoes and improving the phosphorus availability in soils. Future studies should further explore the specific pathways and molecular mechanisms through which oxygen-mycorrhiza synergy influence soil phosphorus availability, integrating root exudates and microbial community data within the "soil oxygen environment-microorganisms-plant physiology" framework. Declarations Availability of data and materials: The data that supports the findings of this study are available from the corresponding author upon reasonable request. Author contributions: Yingji Lian : Investigation, Data curation, Writing – original draft, Visualization; Hongjun Lei : Conceptualization, Resources, Writing – review & editing, Supervision, Project administration, Funding acquisition; Hongwei Pan : Writing – review & editing, Resources; Muhammad Zain : Language polishing, Review & editing; Xin Liu : Software support , Visualization; Shaobo Wang: Writing – review & editing; Yong Liu : Experimental instruction, Supervision; Shihui Zhang : Data analysis, Investigation. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-7489363","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":509432381,"identity":"a91467d4-3ae2-4409-9e5a-773b284b97f2","order_by":0,"name":"Yingji Lian","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAABE0lEQVRIie3Rv2qDQBzA8QsHl+Ug64mxz/ArB6bD0b6KItyUQMeOSsAupl0t9CF8BMNBs/gAhQ61BJw6OKYgtOYK0uGkGTPcF5z8fbw/ImSznWGwS5I6AEYJnm7r9s+bSTxGKqWgvRXz2TSLLnME/xMnl9LJWymcrPRdegqZoSW4FBSF1yDiouvC2F03vRVeUeKmNhCCKuC/JNzuVymE8fzluJzkRUkWYCKTDKJhlVXcE7YkbxSpsCgpYaaNYT2vie9edQP5HiVOSsIkB0n18REZSDlKoMIKtSCovuRNynnK5OLrGSL+pIhvJO8f94egYzf6Vx66C++RRXv4vLv2HnbrxkRMd6g/1T/4pHmbzWazGfoBiylkmuXBOSIAAAAASUVORK5CYII=","orcid":"https://orcid.org/0000-0003-2562-4349","institution":"North China University of Water Resources and Electric Power","correspondingAuthor":true,"prefix":"","firstName":"Yingji","middleName":"","lastName":"Lian","suffix":""},{"id":509432382,"identity":"57c2d236-ea66-4e66-8f91-0886f5da9d52","order_by":1,"name":"Hongjun Lei","email":"","orcid":"https://orcid.org/0009-0009-0528-339X","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Hongjun","middleName":"","lastName":"Lei","suffix":""},{"id":509432383,"identity":"ef1e697e-42e3-4800-af93-7f3642f41c14","order_by":2,"name":"Hongwei Pan","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Hongwei","middleName":"","lastName":"Pan","suffix":""},{"id":509432384,"identity":"1e37e98a-d16d-4da9-80c7-27693c4db75e","order_by":3,"name":"Muhammad Zain","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Muhammad","middleName":"","lastName":"Zain","suffix":""},{"id":509432387,"identity":"7a3d49cb-61c9-4d29-b678-aeac69484adf","order_by":4,"name":"Xin Liu","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Xin","middleName":"","lastName":"Liu","suffix":""},{"id":509432390,"identity":"7567aa7b-d9f6-4382-9e10-557dd8bf6460","order_by":5,"name":"Shaobo Wang","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Shaobo","middleName":"","lastName":"Wang","suffix":""},{"id":509432391,"identity":"06a4a99f-de68-431f-806b-668ce42a0fef","order_by":6,"name":"Yong Liu","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Yong","middleName":"","lastName":"Liu","suffix":""},{"id":509432393,"identity":"c880b47d-9222-4673-bef6-0c958682eace","order_by":7,"name":"Shihui Zhang","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Shihui","middleName":"","lastName":"Zhang","suffix":""}],"badges":[],"createdAt":"2025-08-29 14:12:49","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-7489363/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-7489363/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":90836918,"identity":"fb9707d9-0b8d-41aa-8591-52ec429cc16c","added_by":"auto","created_at":"2025-09-08 17:57:21","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":35283,"visible":true,"origin":"","legend":"\u003cp\u003eMycorrhizal infection rate of tomato under different treatments is presented as percentage of root length colonized by mycorrhizal fungi. Error bars are standard errors (n=3). Small letters above the columns indicate significant differences. C represents standard drip irrigation, and A represents aerated drip irrigation. RI refers to the treatment with \u003cem\u003eRhizophagus intraradices\u003c/em\u003e (RI) inoculation only; FM refers to the treatment with \u003cem\u003eFunneliformis mosseae\u003c/em\u003e (FM) inoculation only; FMRI refers to the mixed inoculation of FM and RI. The same applies to the following text.\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-7489363/v1/6b56119ad08403833e231352.png"},{"id":90836417,"identity":"ebc16b25-97e5-4e79-85b5-c1f5afb543da","added_by":"auto","created_at":"2025-09-08 17:49:21","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":215882,"visible":true,"origin":"","legend":"\u003cp\u003eRoot morphological characteristics of tomato under different irrigation and mycorrhizal fungi inoculation treatments. Among them, (a) represents total root length; (b) represents root surface area; (c) represents root volume; (d) represents root tip number; (e) represents average root diameter.\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-7489363/v1/23445fbd76cd91149c0a0b23.png"},{"id":90836414,"identity":"40d63010-bd41-44c6-8a79-15ee4c09490e","added_by":"auto","created_at":"2025-09-08 17:49:21","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":60856,"visible":true,"origin":"","legend":"\u003cp\u003eEffects of different irrigation modes and mycorrhizal fungi treatments on easily available P fractions in soil.\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-7489363/v1/5d7eaaf5bfd97eba2f4b783f.png"},{"id":90836415,"identity":"15b6dae3-1825-4635-877f-567ec0d6865c","added_by":"auto","created_at":"2025-09-08 17:49:21","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":45667,"visible":true,"origin":"","legend":"\u003cp\u003eEffects of different irrigation modes and mycorrhizal fungi treatments on moderately available P fractions in soil.\u003c/p\u003e","description":"","filename":"4.png","url":"https://assets-eu.researchsquare.com/files/rs-7489363/v1/cc6a5f18ae8a55673476fae7.png"},{"id":90836921,"identity":"b906d88d-48ce-4bb0-96eb-5180b6332ed4","added_by":"auto","created_at":"2025-09-08 17:57:21","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":60183,"visible":true,"origin":"","legend":"\u003cp\u003eEffects of different irrigation modes and mycorrhizal fungi treatments on insoluble P components in soil.\u003c/p\u003e","description":"","filename":"5.png","url":"https://assets-eu.researchsquare.com/files/rs-7489363/v1/d41799eac78639f4e8f70195.png"},{"id":90836422,"identity":"6edd8c55-fd5c-4490-b6a1-9a826136e80c","added_by":"auto","created_at":"2025-09-08 17:49:21","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":35327,"visible":true,"origin":"","legend":"\u003cp\u003eEffects of different irrigation and mycorrhizal fungi treatments on soil ACP activity in tomato.\u003c/p\u003e","description":"","filename":"6.png","url":"https://assets-eu.researchsquare.com/files/rs-7489363/v1/a642b774e2ffbea96da4c20d.png"},{"id":90836419,"identity":"827e1787-0e67-4813-b5c3-7d95a7857d0f","added_by":"auto","created_at":"2025-09-08 17:49:21","extension":"png","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":76201,"visible":true,"origin":"","legend":"\u003cp\u003eEffects of oxygen-mycorrhiza synergy on phosphorus uptake in greenhouse tomato.\u003c/p\u003e","description":"","filename":"7.png","url":"https://assets-eu.researchsquare.com/files/rs-7489363/v1/735c0a8089fd3970368f4d6c.png"},{"id":90838013,"identity":"2b829edf-aa1b-4a40-9c6f-c3a75c80b660","added_by":"auto","created_at":"2025-09-08 18:21:21","extension":"png","order_by":8,"title":"Figure 8","display":"","copyAsset":false,"role":"figure","size":192796,"visible":true,"origin":"","legend":"\u003cp\u003eSEM analysis of the effects of ADI and AMF on tomato yield and phosphorus accumulation. The structural equation model shows the relationships between soil phosphatase activity, phosphorus fractions, tomato root morphology, crop phosphorus accumulation, and yield variables under the synergistic effect of ADI and AMF. Red and blue arrows represent positive and negative relationships, respectively. The numbers near the path arrows represent path coefficients. Solid and dashed lines represent significant and non-significant relationships, with P values indicating significance at the levels\u003cem\u003e of p \u003c/em\u003e= 0.05, 0.01, and 0.001, respectively. R\u003csup\u003e2\u003c/sup\u003e represents the proportion of variance explained by each dependent variable. Resin-exchangeable phosphorus (Resin-P), NaHCO\u003csub\u003e3\u003c/sub\u003e-Pi, and C-HCl-Pi were combined into a composite variable for soil inorganic phosphorus. NaHCO\u003csub\u003e3\u003c/sub\u003e-Po, NaOH-Po, D-HCl-P, and C-HCl-Po were combined into a composite variable for soil organic phosphorus. Tomato root length, root surface area, root volume, root tip number, and mycorrhizal colonization rate were combined into a composite variable for root morphology development. The phosphorus accumulation in tomato stems, leaves, and fruit was combined into a composite variable for plant phosphorus accumulation.\u003c/p\u003e","description":"","filename":"8.png","url":"https://assets-eu.researchsquare.com/files/rs-7489363/v1/cbfd7b620766b2751c324968.png"},{"id":100373336,"identity":"9f018b79-c735-4193-97ba-aa8dc94b626d","added_by":"auto","created_at":"2026-01-16 08:14:05","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2277157,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7489363/v1/abc78858-805b-4a47-90ea-c69d804fd797.pdf"}],"financialInterests":"","formattedTitle":"Oxygen-mycorrhiza synergy drives phosphorus transformation in soil and its accumulation in greenhouse-grown tomato.","fulltext":[{"header":"1. Introduction","content":"\u003cp\u003ePhosphorus is an essential macronutrient for plant growth and development, playing a critical role in energy transfer, photosynthesis, nutrient transport, and metabolic processes. It is particularly important for root development, flowering, and fruit formation in tomato (\u003cem\u003eSolanum lycopersicum L.\u003c/em\u003e), directly affecting both yield and quality (Khan et al. 2023). However, in controlled agricultural systems, due to high crop rotation indices and excessive fertilization, phosphorus in the soil is easily fixed by iron, aluminum oxides, or calcium salts, forming insoluble phosphates. As a result, the proportion of available phosphorus to plants is typically less than 20% (Zhang et al. 2019; Fang et al. 2023; N\u0026eacute;grel et al. 2024). The low phosphorus use efficiency in controlled systems is a critical bottleneck limiting the high yield and sustainability of greenhouse tomato production. Traditional strategies rely on increasing the application of phosphorus fertilizers, which not only raises production costs but also leads to significant accumulation of residual phosphorus in the soil (Pavinato et al. 2020), which ultimately results in soil degradation and environmental issues. Therefore, there is an urgent need to explore innovative technologies that can effectively activate fixed phosphorus in the soil and improve plant phosphorus absorption and utilization efficiency (Song et al. 2020).\u003c/p\u003e\u003cp\u003eResearchers have actively explored solutions to address the issue of phosphorus fixation. Arbuscular mycorrhizal fungi (AMF) are soil microorganisms closely related to plant nutrition (Tedersoo et al. 2020) that form symbiotic relationships with approximately 80% of terrestrial plants (Branco et al. 2022; Shi et al. 2023). These AMF are reported to be involved in significantly extending the root absorption range and enhancing the uptake of water and nutrients (Kuila and Ghosh 2022), particularly phosphorus (Peng et al. 2020). The hyphal network of AMF can directly absorb phosphates from the soil and transfer them to the plant roots through the mycorrhizal interface. Additionally, AMF can promote the mineralization of organic phosphorus by secreting acid phosphatases (Chen et al. 2020). Aerated drip irrigation (ADI), an advanced version of subsurface drip irrigation, improves the soil dissolved oxygen content, which enhances the rhizosphere microecological environment, increases soil oxygen levels, and the air-filled porosity (Ouyang et al. 2025; Wang et al. 2024). This in turn boosts microbial activity, promotes root growth, and facilitates nutrient transformation. Dong et al. (2025) found that ADI significantly increased root length by 19.02%, root volume by 22.92%, and root vitality by 32.1% as compared to traditional drip irrigation. Furthermore, ADI can indirectly promote nutrient cycling by altering soil microbial community structure and increasing the abundance of beneficial microorganisms (Zhao et al. 2019; Zhu et al. 2022). Research on vertical flow constructed wetlands has shown that intermittent aeration can increase the AMF colonization rate to 67% (Xu et al. 2021), indicating that moderate aeration can optimize AMF community structure.\u003c/p\u003e\u003cp\u003eAlthough both ADI and AMF have shown progress in improving phosphorus use efficiency when applied individually, but still there is limited systematic research on their synergistic effects. The environmental factors such as high temperature, high humidity, and low oxygen levels may weaken AMF colonization efficiency and function, specifically in controlled soils (Carvalho et al. 2015; Huo et al. 2021). In addition, the application of ADI alone may not fully exert its regulatory potential on soil microbial communities, limiting its effectiveness in enhancing phosphorus utilization and plant stress resistance (Cao et al. 2024; Chen et al. 2023). The research gap hinders the development of strategies to optimize phosphorus management in controlled agricultural systems. Therefore, we hypothesized that comprehensive study on the synergistic effects of ADI and AMF on soil phosphorus transformation, root morphology, and phosphorus allocation, as well as the interactive effects of different AMF strains (such as \u003cem\u003eRhizophagus intraradices\u003c/em\u003e and \u003cem\u003eFunneliformis mosseae\u003c/em\u003e) under oxygenated conditions should be conducted. We planned a study with objectives to (\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e) investigates the synergistic effects of ADI and AMF through a two-factor experiment (irrigation mode \u0026times; AMF inoculation) and systematically analyzes the changes in Hedley soil phosphorus fractions, root morphology, and soil phosphatase activity in response to oxygen-mycorrhiza synergy; (\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e) quantify the contribution of AMF colonization rate and soil phosphorus fraction changes to phosphorus uptake and yield in tomato plants through Structural equation modeling (SEM). The findings from this study will provide new insights into addressing phosphorus fixation in controlled soils and improving phosphorus fertilizer use efficiency.\u003c/p\u003e"},{"header":"2. Materials and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e\u003ch2\u003e2.1 Experimental materials\u003c/h2\u003e\u003cp\u003eThe tested tomato variety \u003cem\u003eSolanum lycopersicum\u003c/em\u003e \"Huafei No. 1\", was acquired from the Xinxiang Breeding Base. Two types of arbuscular mycorrhizal fungi (AMF) inoculants were used: one was \u003cem\u003eFunneliformis mosseae\u003c/em\u003e (FM), purchased from the China AMF Resource Bank (BGC, XJ01) at the Beijing Academy of Agriculture and Forestry Sciences. The spore concentration was approximately 230 spores g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, including the corresponding culture medium, spores, mycelium, and plant root segments. The other inoculant \u003cem\u003eRhizophagus intraradices\u003c/em\u003e (RI) was taken from Foshan Ironman Environmental Technology Co., Ltd. The inoculant contained fine sand, spores, mycelium, and plant root segments, with a spore concentration of approximately 70 spores g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e.\u003c/p\u003e\u003cp\u003eThe experiment was conducted in pots filled with clay soil. The pots were placed in a modern greenhouse at the Agricultural Efficient Water Use Experimental Field of North China University of Water Resources and Electric Power. The greenhouse was equipped with wet curtains, fans, and shading nets to regulate temperature and humidity. The soil was sterilized by high-temperature treatment: it was first sieved through a 2 mm mesh, then placed in an oven set to 160\u0026deg;C for 2 hours. After cooling, the sterilization process was repeated for another 2 hours. The soil was left to cool naturally before use (Hu et al. 2020). The basic chemical properties of the soil are given in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eBasic chemical properties of tested soils.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"9\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSoil type\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eBulk density\u003c/p\u003e\u003cp\u003e(g\u0026middot;cm\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eField capacity\u003c/p\u003e\u003cp\u003e(%)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eOrganic matter\u003c/p\u003e\u003cp\u003e(g\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eNitrate nitrogen\u003c/p\u003e\u003cp\u003e(mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAmmonium nitrogen\u003c/p\u003e\u003cp\u003e(mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003eTotal phosphorus\u003c/p\u003e\u003cp\u003e(g\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAvailable phosphorus\u003c/p\u003e\u003cp\u003e(mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c9\"\u003e\u003cp\u003epH\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eClay\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e1.32\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e25.94\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e11.62\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e21.4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e4.55\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e58.43\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e8.11\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec4\" class=\"Section2\"\u003e\u003ch2\u003e2.2 Experimental design\u003c/h2\u003e\u003cp\u003eThe experiment was a two-factor fully factorial design with the following treatments. (\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e) Irrigation Modes: (a) Conventional standard drip irrigation without additional aeration (C) has a dissolved oxygen (DO) value of approximately 6 mg\u0026middot;L\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e in the irrigation water; (b) Aerated drip irrigation (A) with a DO value set to 15 mg\u0026middot;L\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e in the irrigation water; (\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e) AMF Inoculation Treatments: (a) \u003cem\u003eFunneliformis mosseae\u003c/em\u003e (FM) inoculation only; (b) \u003cem\u003eRhizophagus intraradices\u003c/em\u003e (RI) inoculation only; (c) A mixed inoculation with both fungi, each at 50% of the individual spore concentration (FMRI). Based on the spore concentration of the inoculants, the FM treatment received 5g of \u003cem\u003eFunneliformis mosseae\u003c/em\u003e, the RI treatment received 16g of \u003cem\u003eRhizophagus intraradices\u003c/em\u003e, and the FMRI treatment received 2.5g of \u003cem\u003eFunneliformis mosseae\u003c/em\u003e and 8g of \u003cem\u003eRhizophagus intraradices\u003c/em\u003e. This resulted in a total of 8 treatments, with 5 replications per treatment, totaling 40 pots. The details of experimental design are presented in Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eExperimental treatments with their respective DO and inoculants concentrations.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"4\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eTreatment\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eDO in irrigation water\u003c/p\u003e\u003cp\u003e(mg\u0026middot;L\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cem\u003eFunneliformis mosseae\u003c/em\u003e\u003c/p\u003e\u003cp\u003e(g\u0026middot;pot\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eRhizophagus intraradices\u003c/em\u003e\u003c/p\u003e\u003cp\u003e(g\u0026middot;pot\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCRI\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e16\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCFM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e5.0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCFMRI\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eA\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e15\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eARI\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e15\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e16\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eAFM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e15\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e5.0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eAFMRI\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e15\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eThe numbers in the table represent the levels of different independent variables. DO represents the dissolved oxygen concentration of irrigation water; C represents standard drip irrigation without additional aeration; A represents aerated drip irrigation; RI refers to the treatment with \u003cem\u003eRhizophagus intraradices\u003c/em\u003e (RI) inoculation only; FM refers to the treatment with \u003cem\u003eFunneliformis mosseae\u003c/em\u003e (FM) inoculation only; FMRI refers to the mixed inoculation of FM and RI.\u003c/p\u003e\u003cp\u003eThe pots used for the experiment were 22 cm in diameter and 27 cm in height. After sterilizing the soil with high-temperature treatment, each pot was filled with 8 kg of dry soil, mixed with 2.0 g of calcium superphosphate as a basal fertilizer. This mixture was irrigated with 500 mL of water on the same day, and the soil was left to settle and infiltrate overnight, followed by supplemental watering based on the soil moisture condition on the next day until the soil moisture content reached approximately 70% of the field capacity, at which point transplanting was conducted. A hole (3\u0026ndash;5 cm deep) was dug in each pot, arbuscular mycorrhizal fungi (AMF) were placed in the hole in a flat layer, and then one tomato seedling with 4 true leaves and 1 heart or 5 true leaves and 1 heart was transplanted into the pot, followed by compaction of the surrounding soil. After transplantation, a soil solution was prepared by mixing the soil with deionized water at a 2:1 soil-to-water ratio, and the mixture was wet-sieved through a 30 \u0026micro;m filter to remove large soil microorganisms, including AMF spores. The filtrate was then quantified (20 ml\u0026middot;pot\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) and added to the pots to restore the basic microbial community of the sterilized soil (Sun et al. 2022).\u003c/p\u003e\u003cp\u003e\u003cb\u003eFertilization management\u003c/b\u003e: Each pot was fertilized with urea (46% N content), calcium superphosphate (12% P content), and potassium sulfate (50% K content) as nitrogen, phosphorus, and potassium fertilizers, respectively. Corresponding to the following field rates: 180 kg nitrogen\u0026middot;ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, 90 kg phosphorus\u0026middot;ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, and 240 kg potassium\u0026middot;ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, each pot received a total of 1.57 g urea, 1.92 g potassium sulfate, and 3.0 g calcium superphosphate during the growing season. The calcium superphosphate was applied as a basal fertilizer, while urea and potassium sulfate were applied in a ratio of 1:3:2 at the seedling, flowering, and early fruiting stages, respectively, through fertigation.\u003c/p\u003e\u003cp\u003e\u003cstrong\u003eIrrigation method\u003c/strong\u003e\u003cp\u003eThe irrigation system consists of buried drip emitters, with an emitter working pressure of 0.10 Mpa. Standard drip irrigation treatments received water through the standard supply system, while for aerated treatments we utilized a Venturi air injector (Mazzei air injector 684, Mazzei Corp. USA) to provide oxygenation. Soil moisture content was monitored using a handheld Time Domain Reflectometry (TDR) device. When the soil moisture content dropped to approximately 60% of field capacity, irrigation was performed on the base of evaporation rate from a standard evaporation pan. The amount of irrigation water was calculated using the following Eq.\u0026nbsp;(\u003cspan refid=\"Equ1\" class=\"InternalRef\"\u003e1\u003c/span\u003e)\u003c/p\u003e\u003c/p\u003e\u003cp\u003e\u003cdiv id=\"Equ1\" class=\"Equation\"\u003e\u003cdiv format=\"TEX\" class=\"mathdisplay\" id=\"FileID_Equ1\" name=\"EquationSource\"\u003e\n$$W=A \\cdot {E_P} \\cdot {K_P}$$\u003c/div\u003e\u003cdiv class=\"EquationNumber\"\u003e1\u003c/div\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eWhere \u003cem\u003eW\u003c/em\u003e\u0026mdash;Single irrigation amount (L); \u003cem\u003eA\u003c/em\u003e\u0026mdash;Area of the planting pot (m\u003csup\u003e2\u003c/sup\u003e) (which was 0.038 m\u003csup\u003e2\u003c/sup\u003e in this study); \u003cem\u003eE\u003c/em\u003e\u003csub\u003e\u003cem\u003eP\u003c/em\u003e\u003c/sub\u003e\u0026mdash;Evaporation from the pan between two irrigation events (mm); \u003cem\u003eK\u003c/em\u003e\u003csub\u003e\u003cem\u003eP\u003c/em\u003e\u003c/sub\u003e\u0026mdash;Pan coefficient, taken as 0.8.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec5\" class=\"Section2\"\u003e\u003ch2\u003e2.3 Sample collection and measurement\u003c/h2\u003e\u003cp\u003eAfter the tomatoes matured on January 21st, 2025, the plants were carefully removed from the soil. The soil samples were air-dried naturally and used to determine the soil physicochemical properties and phosphatase activity. The tomato plants were washed with deionized water and then separated into aboveground and underground parts. The fresh roots were scanned, weighed, and then a portion of about 1 cm of root segments was collected to determine the mycorrhizal colonization rate. After weighing the fresh root mass, the samples were blanched at 105\u0026deg;C for 30 minutes, followed by drying at 60\u0026deg;C to a constant weight. The dry weight was recorded, and the samples were then ground and passed through a 0.149 mm sieve. The phosphorus contents in both aboveground and underground parts of the plant were measured using the H\u003csub\u003e2\u003c/sub\u003eSO\u003csub\u003e4\u003c/sub\u003e-H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e digestion method and molybdenum-antimony colorimetry. The phosphorus accumulation was calculated, and the phosphorus content in each part was determined using an AA3 Flow Analyzer. Rhizosphere soil samples were collected to measure available phosphorus and total phosphorus.\u003c/p\u003e\u003cp\u003e\u003cb\u003e2.3.1. Mycorrhizal colonization rate\u003c/b\u003e: The root staining method was consists of improved acetic acid ink staining technique (Vierheilig et al. 1998). The colonization rate was calculated using the method widely adopted by the Arbuscular Mycorrhizal Fungi Germplasm Resource Bank (BGC), which determines the overall colonization rate as described by Biermann and Linderman (1981):\u003cdiv id=\"Equ2\" class=\"Equation\"\u003e\u003cdiv format=\"TEX\" class=\"mathdisplay\" id=\"FileID_Equ2\" name=\"EquationSource\"\u003e\n$${\\text{Mycorrhizal Colonization Rate}}(\\% )=\\frac{{\\sum {(0 \\times {S_i}+10 \\times {S_i}+20 \\times {S_i}+ \\cdots 100 \\times {S_i})} }}{{{\\text{Total EquationNumber of root segments observed}}}}$$\u003c/div\u003e\u003cdiv class=\"EquationNumber\"\u003e2\u003c/div\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eUnder the microscope, the mycorrhizal colonization status of each root segment was examined (e.g., points of penetration, hyphae, arbuscules, vesicles, etc.). The colonization rate was scored according to the infection percentage, with values of 0, 10, 20, 30, \u0026hellip;, 100, where 10 represents the first level of infection. For each segment, the number of root segments infected at each level (Si) was recorded. The mycorrhizal colonization rate of the sample was calculated by dividing the sum of product of the number of root segments and their respective levels by the total number of observed segments (Eq.\u0026nbsp;\u003cspan refid=\"Equ2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003cb\u003e2.3.2 Soil phosphorus fractionations\u003c/b\u003e: The phosphorus fractionations in the soil were determined using the modified Hedley phosphorus fractionation method (Tiessen and Moir, 1993). In brief, 0.5 g of air-dried soil passed through a 100-mesh sieve was weighed and treated with 30 mL deionized water (containing anion exchange resin) to extract water-soluble phosphorus (Resin-Pi) using the molybdenum-antimony colorimetric method. Subsequently, the soil was sequentially extracted with 0.5 M NaHCO\u003csub\u003e3\u003c/sub\u003e, 0.1 M NaOH, and 1.0 M HCl (D-HCl-Pi), with the C-HCl-P fraction extracted by heating in a water bath. Residual phosphorus (Residual-P) was extracted using H\u003csub\u003e2\u003c/sub\u003eSO\u003csub\u003e4\u003c/sub\u003e-H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e digestion. Finally, the fractions were quantified using the molybdenum-antimony colorimetric method.\u003c/p\u003e\u003cp\u003e\u003cb\u003e2.3.3 Soil phosphatase activity\u003c/b\u003e: Soil acid phosphatase activity was measured using the p-nitrophenyl phosphate disodium (PNPP) colorimetric method (Veloso et al. 2023; Tabatabai and Bremner, 1969). Accurately weighed 0.20 g of soil was passed through a 0.18 mm sieve and placed into a 10 mL round-bottom plastic centrifuge tube. Later on, 0.2 mL toluene was added, vortex to mix, and left to stand for 15 minutes. Then, 4 mL phosphate buffer and 1 mL of 0.05 M p-nitrophenyl phosphate disodium solution (prepared in phosphate buffer) was added. After capping, it was mixed thoroughly and incubated at 37\u0026deg;C for 1 hour. After incubation, shake the mixture, and added 1 mL of 0.5 M CaCl\u003csub\u003e2\u003c/sub\u003e and 8 mL of 0.5 M NaOH, vortex to mix, and centrifuge at 2500 rpm for 5 minutes. Next, we transferred 5 mL of the supernatant to a new tube and centrifuge at 3800 rpm for 5 minutes. Finally, we measured the absorbance at 410 nm. The acid phosphatase activity in the soil is expressed as the micrograms of p-nitrophenol per gram of soil per unit time, with units of mg\u0026middot;g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u0026middot;d\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e.\u003c/p\u003e\u003cp\u003e\u003cb\u003e2.3.4 Root morphology parameters\u003c/b\u003e: After washing the roots with deionized water, the clean roots were placed in a transparent tray with a depth of 10 mm. Using forceps, the roots were gently spread out to ensure full extension. The roots were then scanned using a root scanner (Epson Expression 1600 Pro, Japan). The scanned root images were analyzed with the WinRHIZO image analysis system (Regent, Canada) to measure root morphology parameters such as total root length, root volume, and root surface area.\u003c/p\u003e\u003cdiv id=\"Sec6\" class=\"Section3\"\u003e\u003ch2\u003e\u003cb\u003e2.3.5 Crop phosphorus accumulation and phosphorus use efficiency\u003c/b\u003e:\u003c/h2\u003e\u003cp\u003e(\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e) \u003cb\u003ePhosphorus accumulation in plants\u003c/b\u003e: The root, stem, leaf, and fruit samples were washed with clean water, then placed in an oven at 105\u0026deg;C for 15 minutes for blanching. Afterward, the samples were dried at 75\u0026deg;C to a constant weight and weighed. After weighing, the samples were ground through a 0.15 mm-mesh sieve and stored in bags for further analysis. The total phosphorus content in each organ was determined using the H\u003csub\u003e2\u003c/sub\u003eSO\u003csub\u003e4\u003c/sub\u003e-H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e digestion method, and phosphorus accumulation in each organ was calculated using a flow analyzer (AA3, SEAL Analytical, Germany).\u003c/p\u003e\u003cp\u003e(\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e) \u003cb\u003eMycorrhizal growth response (MGR) and mycorrhizal phosphorus uptake response (MPR)\u003c/b\u003e: The \u003cem\u003eMGR\u003c/em\u003e and \u003cem\u003eMPR\u003c/em\u003e were used to assess the impact of AMF symbiosis on tomato growth and phosphorus uptake in each treatment (Veiga et al. 2011). The \u003cem\u003eMGR\u003c/em\u003e was calculated as follows:\u003cdiv id=\"Equ3\" class=\"Equation\"\u003e\u003cdiv format=\"TEX\" class=\"mathdisplay\" id=\"FileID_Equ3\" name=\"EquationSource\"\u003e\n$$N{M_{mean}}\u0026lt;AM,{\\text{ }}MGR=\\left( {1 - \\frac{{N{M_{mean}}}}{{AM}}} \\right) \\times 100$$\u003c/div\u003e\u003cdiv class=\"EquationNumber\"\u003e3\u003c/div\u003e\u003c/div\u003e\u003cdiv id=\"Equ4\" class=\"Equation\"\u003e\u003cdiv format=\"TEX\" class=\"mathdisplay\" id=\"FileID_Equ4\" name=\"EquationSource\"\u003e\n$$N{M_{mean}}\u0026gt;AM,{\\text{ }}MGR=\\left( {\\frac{{AM}}{{N{M_{mean}}}} - 1} \\right) \\times 100$$\u003c/div\u003e\u003cdiv class=\"EquationNumber\"\u003e4\u003c/div\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eWhere \u003cem\u003eNM\u003c/em\u003e\u003csub\u003e\u003cem\u003emean\u003c/em\u003e\u003c/sub\u003e represents the average biomass of tomato plants in the non-AMF treatment group, and \u003cem\u003eAM\u003c/em\u003e is the biomass of the tomato plants in each AMF treatment group. The same method applies for calculating \u003cem\u003eMPR\u003c/em\u003e.\u003c/p\u003e\u003cp\u003e(\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e) \u003cb\u003ePhosphorus acquisition efficiency (PAE) and Phosphorus use efficiency (PUE)\u003c/b\u003e: \u003cem\u003ePAE\u003c/em\u003e and \u003cem\u003ePUE\u003c/em\u003e were used to characterize the phosphorus efficiency of the tested tomato plants (Deng et al. 2018; Han et al. 2022). The calculation formulas are as follows:\u003cdiv id=\"Equ5\" class=\"Equation\"\u003e\u003cdiv format=\"TEX\" class=\"mathdisplay\" id=\"FileID_Equ5\" name=\"EquationSource\"\u003e\n$$PAE=\\frac{{{P_{plant}}}}{{{W_{root}}}}$$\u003c/div\u003e\u003cdiv class=\"EquationNumber\"\u003e5\u003c/div\u003e\u003c/div\u003e\u003cdiv id=\"Equ6\" class=\"Equation\"\u003e\u003cdiv format=\"TEX\" class=\"mathdisplay\" id=\"FileID_Equ6\" name=\"EquationSource\"\u003e\n$$PUE=\\frac{Y}{{{P_{\\text{a}}}}}$$\u003c/div\u003e\u003cdiv class=\"EquationNumber\"\u003e6\u003c/div\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eWhere, \u003cem\u003ePAE\u003c/em\u003e represents the ability of the plant to acquire phosphorus from the soil, in mg\u0026middot;g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e; \u003cem\u003eP\u003c/em\u003e\u003csub\u003e\u003cem\u003eplant\u003c/em\u003e\u003c/sub\u003e represents the total phosphorus uptake by the plant in mg\u0026middot;pot\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e; \u003cem\u003eW\u003c/em\u003e\u003csub\u003e\u003cem\u003eroot\u003c/em\u003e\u003c/sub\u003e represents the dry weight of the roots in g\u0026middot;pot\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e. While, \u003cem\u003ePUE\u003c/em\u003e represents the ability of the plant to utilize the acquired phosphorus to generate biomass or yield in kg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e; \u003cem\u003eY\u003c/em\u003e represents the tomato yield in kg\u0026middot;ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e; \u003cem\u003eP\u003c/em\u003e\u003csub\u003e\u003cem\u003ea\u003c/em\u003e\u003c/sub\u003e represents the total phosphorus absorbed by plants per hectare in kg\u0026middot;ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e.\u003c/p\u003e\u003c/div\u003e\u003c/div\u003e\u003cdiv id=\"Sec7\" class=\"Section2\"\u003e\u003ch2\u003e2.4 Data analysis\u003c/h2\u003e\u003cp\u003ePrior to data analysis, outliers were removed, and the data were tested for normality and homogeneity of variance. A multivariate analysis of variance (Multivariate ANOVA) was conducted to examine the individual and interactive effects of irrigation water DO and AMF inoculation modes on tomato root morphology, yield, phosphorus accumulation, and soil phosphorus fractions. Duncan's post-hoc multiple comparisons were performed for significance testing, with a significance level set at 0.05. All data were analyzed using SPSS 25.0 (IBM, Armonk, NY, USA) statistical software. Structural equation modeling (SEM) was constructed in SmartPLS 4.1.1.4 (SmartPLS, Hamburg, Germany) to evaluate the direct and indirect effects of \"fungal colonization rate, phosphatase activity, and root morphology\" on soil phosphorus activation and tomato phosphorus utilization. Graphs were generated using Origin 2021 software (OriginLab Corporation, Northampton, MA, USA).\u003c/p\u003e\u003c/div\u003e"},{"header":"3. Results and analysis","content":"\u003cdiv id=\"Sec9\" class=\"Section2\"\u003e\u003ch2\u003e3.1 Arbuscular mycorrhizal fungi colonization\u003c/h2\u003e\u003cp\u003eAerated drip irrigation (ADI) significantly affected the mycorrhizal colonization rate of arbuscular mycorrhizal fungi (AMF) in greenhouse grown tomatoes (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05) (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The results revealed that non-inoculated treatments (C and A) had 0% colonization rate, indicating that the soil sterilization was effective prior to the experiment and that the system was free from background AMF contamination. Under standard drip irrigation condition (C), the highest colonization rate (20.32%) was observed in the \u003cem\u003eRhizophagus intraradices\u003c/em\u003e (RI) inoculation treatment, which was 14.86% higher than the inoculation of \u003cem\u003eFunneliformis mosseae\u003c/em\u003e (FM) treatment, while the mixed inoculation (CFMRI) treatment had the lowest (14.68%) colonization rate. Under aerated conditions, the ARI treatment had the highest colonization rate of 26.95%, which was 32.66% higher than the CRI treatment and represented the highest rate across all treatments. The next highest inoculation rate (20.60%) was observed in AFM treatment followed by the AFMRI treatment (19.44%).\u003c/p\u003e\u003cp\u003eOverall, the RI inoculation treatment exhibited a higher colonization rate under both irrigation conditions, while no significant difference was found between the RI and RIFM mixed inoculation treatments. This suggests that the RI inoculant was more effective at colonizing tomatoes in this system compared to FM, which may be attributed to differences in host adaptation between the strains. Additionally, the colonization rate of mixed inoculation treatment is lower than that of RI inoculation treatment alone, indicating that there may be competition or inhibition effect between mixed inoculation agents. Furthermore, the mycorrhizal colonization rate under ADI conditions was generally higher than that of standard drip irrigation, which may be related to the improved soil aeration by ADI, facilitating the germination and infection of fungal spores. The ADI system regulation of the rhizosphere oxygen environment, enhancing AMF activity, may further regulate phosphorus transformation and utilization, as well as tomato yield formation. This combined use of ADI and AMF and their effect is defined as the \"oxygen-mycorrhiza synergy\".\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec10\" class=\"Section2\"\u003e\u003ch2\u003e3.2 Root morphology characteristics\u003c/h2\u003e\u003cp\u003eThe results of root morphology parameters under different treatments are given in Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e. It was observed that, ADI significantly upregulated the total root length, root surface area, and root tip number in tomatoes (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05). The overall trends of total root length, root surface area, root volume, and root tip number were consistent, with the ARI treatment showing the highest values, the C treatment without inoculation showing the lowest. While, A treatment without inoculation and the CRI treatment with single inoculation were falling between these two high and low extremes (ARI\u0026thinsp;\u0026gt;\u0026thinsp;A\u0026thinsp;\u0026asymp;\u0026thinsp;CRI\u0026thinsp;\u0026gt;\u0026thinsp;C). No significant patterns were observed in the average root diameter, and neither ADI nor AMF treatments had a significant impact on this parameter.\u003c/p\u003e\u003cp\u003eUnder standard drip irrigation conditions (C), inoculation with \u003cem\u003eRhizophagus intraradices\u003c/em\u003e (CRI) significantly improved root morphology: total root length (980.02 cm), root surface area (383.82 cm\u003csup\u003e2\u003c/sup\u003e), root volume (11.97 cm\u003csup\u003e3\u003c/sup\u003e), and root tip number (2194) which were increased by 13.24%, 17.51%, 21.83%, and 13.25%, respectively, as compared to their control treatment (C). Inoculation with \u003cem\u003eFunneliformis mosseae\u003c/em\u003e (CFM) also showed some enhancement in these four root parameters (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;0.05), but the increases were less than those observed with the RI inoculation. The mixed inoculation (CFMRI) treatment did not demonstrate any synergistic advantage; its root length, root surface area, and root tip number were similar to those of the C treatment without inoculation. However, in terms of average root diameter, the mixed inoculation treatment showed the highest value of 1.27 mm, which was a 5.85% increase compared to the control (C).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eADI enhances the positive effects of AMF inoculation. Except for the average root diameter, the root morphology parameters under the RI inoculation treatment (ARI) in aerated conditions were the highest across all treatments: total root length (1211.82 cm) was 23.65% higher than the standard drip irrigation CRI treatment; root surface area (460.42 cm\u003csup\u003e2\u003c/sup\u003e) increased by 19.96%; root volume (14.11 cm\u003csup\u003e3\u003c/sup\u003e) increased by 17.90%; and the number of root tips (2714) increased by 23.70% compared to CRI. Under aerated conditions, inoculation with \u003cem\u003eFunneliformis mosseae\u003c/em\u003e (FM) and the mixed inoculation (FMRI) also showed improvements in root length, surface area, and root tip number compared to standard drip irrigation, but the increases were less significant than those observed with RI inoculation. Furthermore, the mixed inoculation treatment did not perform well under both irrigation modes except for the average root diameter being similar to or even slightly lower than those under FM treatment.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec11\" class=\"Section2\"\u003e\u003ch2\u003e3.3 Soil phosphorus fractionations and soil phosphatase activity\u003c/h2\u003e\u003cp\u003eThe modified Hedley phosphorus fractionation method (Tiessen and Moir 1993) divides soil phosphorus into 9 fractions. In this study, we classified soil phosphorus into three categories based on its ease of absorption by plants: (\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e) Easily available phosphorus, which includes resin-exchangeable phosphorus (Resin-P), and two NaHCO\u003csub\u003e3\u003c/sub\u003e-extracted phosphorus fractions (NaHCO\u003csub\u003e3\u003c/sub\u003e-Pi, NaHCO\u003csub\u003e3\u003c/sub\u003e-Po); (\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e) Moderately available phosphorus, which includes two NaOH-extracted phosphorus fractions (NaOH-Pi, NaOH-Po); (\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e) Insoluble phosphorus, which includes phosphorus extracted with diluted hydrochloric acid (D-HCl-P), two phosphorus fractions extracted with concentrated hydrochloric acid (C-HCl-Pi, C- HCl-Po), and residual phosphorus (Residual-P). The changes in each phosphorus fraction under different treatments are summarized as follows.\u003c/p\u003e\u003cdiv id=\"Sec12\" class=\"Section3\"\u003e\u003ch2\u003e3.3.1 Easily available phosphorus (Easily available P)\u003c/h2\u003e\u003cp\u003eThe results for directly and easily available phosphorous for plant uptake or rapidly mineralized into available phosphorus including Resin-exchangeable phosphorus (Resin-P), NaHCO\u003csub\u003e3\u003c/sub\u003e-extracted inorganic phosphorus (NaHCO\u003csub\u003e3\u003c/sub\u003e-Pi), and some NaHCO\u003csub\u003e3\u003c/sub\u003e-extracted organic phosphorus (NaHCO\u003csub\u003e3\u003c/sub\u003e-Po) are given in in Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e.\u003c/p\u003e\u003cp\u003eWe found that aeration without inoculation significantly increased (42.03%) the Resin-P content in soil (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.01), showing a value of 18.99 mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e as compared to the control group (C) without inoculation having a value of 13.37 mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e. The CRI, CFM, and mixed inoculation (CFMRI) treatments increased Resin-P by 53.52%, 51.66%, and 25.83%, respectively than that of irrigation without aeration and inoculation (C). On the other hand, ARI, AFM, and AFMRI treatments increased Resin-P by 42.78%, 26.63%, and 22.14%, respectively, as opposed to A treatment without inoculation. Inoculation with RI, FM, and the RI\u0026thinsp;+\u0026thinsp;FM mixed inoculation, combined with ADI, further increased Resin-P by 32.09%, 18.59%, and 37.87%, respectively, compared to the only inoculation treatments (CRI, CFM and CFMRI). In summary, applying arbuscular mycorrhizal inoculation under aerated drip irrigation significantly increased soil Resin-P, and their combined use demonstrated a positive synergistic effect, specifically achieved under combination of ADI and the RI inoculant (ARI).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eNaHCO\u003csub\u003e3\u003c/sub\u003e-Pi is an active inorganic phosphorus extracted by sodium bicarbonate solution, which is mainly in an easily adsorbed form (phosphorus weakly bound to aluminum, iron, and calcium). Compared to the NaHCO\u003csub\u003e3\u003c/sub\u003e-Pi content (37.21 mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) in the C treatment (without inoculation and aeration treatment), both aerated drip irrigation and inoculation with fungi significantly affected the NaHCO\u003csub\u003e3\u003c/sub\u003e-Pi content in the soil (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.01). The value for NaHCO\u003csub\u003e3\u003c/sub\u003e-Pi content in aerated treatment without any inoculation (A) was 44.67 mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, which was 20.04% high compared to control group (C). The CRI, CFM, and mixed inoculation (CFMRI) treatments increased NaHCO\u003csub\u003e3\u003c/sub\u003e-Pi by 82.41%, 57.98%, and 42.39%, respectively, compared to the control group (C). While under aerated irrigation, ARI, AFM, and AFMRI treatments increased NaHCO\u003csub\u003e3\u003c/sub\u003e-Pi by 76.07%, 49.13%, and 43.95%, respectively, compared to aerated treatment (A). When inoculation was done with RI, FM, or the RI\u0026thinsp;+\u0026thinsp;FM under ADI, NaHCO\u003csub\u003e3\u003c/sub\u003e-Pi was further increased by 15.87%, 13.32%, and 21.36%, respectively, compared to CRI, CFM, and CFMRI treatments. Overall, AMF inoculation under aerated drip irrigation increased the soil NaHCO\u003csub\u003e3\u003c/sub\u003e-Pi content, and their combined use demonstrates a significant synergistic effect, with the maximum increase observed in the ADI\u0026thinsp;+\u0026thinsp;RI inoculant combination (ARI).\u003c/p\u003e\u003cp\u003eThe NaHCO\u003csub\u003e3\u003c/sub\u003e-Po is mainly an easily mineralizable organic phosphorus and is an important source of available phosphorus in the soil. In contract to NaHCO\u003csub\u003e3\u003c/sub\u003e-Pi, the value for NaHCO\u003csub\u003e3\u003c/sub\u003e-Po (45.33 mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) in the aerated treatment (A) was 11.01% less as compared to control group value (50.94 mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e). The CRI, CFM, and mixed inoculation (CFMRI) treatments reduced NaHCO\u003csub\u003e3\u003c/sub\u003e-Po by 18.71%, 16.65%, and 7.5%, respectively, compared to the control group (C). The ARI, AFM, and AFMRI treatments reduced NaHCO\u003csub\u003e3\u003c/sub\u003e-Po by 30.25%, 17.36%, and 13.99%, respectively, compared to aerated treatment without inoculation (A). When combined RI, FM, or the RI\u0026thinsp;+\u0026thinsp;FM mixed inoculation with ADI, NaHCO\u003csub\u003e3\u003c/sub\u003e-Po was further reduced by 23.64%, 11.78%, and 17.26%, respectively, compared to inoculation treatments without aeration. Generally, AMF inoculation under aerated drip irrigation led to a reduction in NaHCO\u003csub\u003e3\u003c/sub\u003e-Po, with the maximum reduction observed in the ADI\u0026thinsp;+\u0026thinsp;RI inoculant combination (ARI).\u003c/p\u003e\u003cp\u003eOverall, both ADI and AMF technologies increased the total easily available phosphorus fractions (Resin-P, NaHCO\u003csub\u003e3\u003c/sub\u003e-Pi, and NaHCO\u003csub\u003e3\u003c/sub\u003e-Po) in the soil. Compared to the easily available phosphorus content in the control soil (101.52 mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), aerated drip irrigation (A) had 35.3% increase in this value. Inoculation with AMF also showed similar effects, with RI, FM, and mixed inoculation increasing the total easily available phosphorus by 27.86%, 19.70%, and 15.18%, respectively as compared to CRI, CFM, and CFMRI treatments. Under oxygen-mycorrhiza synergy conditions, this effect was more pronounced, such as ARI, AFM, and AFMRI treatments increased total easily available phosphorus by 35.32%, 26.20%, and 24.59%, respectively, compared to the C treatment. The ADI\u0026thinsp;+\u0026thinsp;RI inoculant (ARI) combination achieved the maximum increase in efficiency.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec13\" class=\"Section3\"\u003e\u003ch2\u003e3.3.2 Moderately available phosphorus (Moderately available P)\u003c/h2\u003e\u003cp\u003eThe NaOH-extracted inorganic phosphorus (NaOH-Pi) and organic phosphorus (NaOH-Po) were categorized as moderately available phosphorus, which lies between easily available phosphorus and insoluble phosphorus. These fractions can be released over a longer time scale through desorption and mineralization. The results obtained from application of irrigation modes and AMF inoculation in tomato plants are presented in Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e. We found that there were no statistical differences among various treatments for NaOH-Pi contents. Generally, the NaOH-Pi value in control group (C treatment) was 84.03 mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, while it was 81.09 mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e in aerated drip irrigation treatment (A), a 3.5% decrease compared to control group (C). The CRI, CFM, and mixed inoculation (CFMRI) treatments reduced NaOH-Pi by 9.13%, 5.95%, and 7.14%, respectively, compared to the control group (C). Further, ARI, AFM, and AFMRI treatments reduced NaOH-Pi by 8.63%, 6.13%, and 2.5%, respectively, that of alone aerated treatment (A). Besides, inoculation with RI, FM in aerated drip irrigation further reduced the NaOH-Pi by 2.98% and 3.69% as opposed to CRI and CFM treatments respectively.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eNaOH-Po is primarily organic phosphorus bound with humus. In contrast to NaOH-Pi, the application of inoculation significantly affected the NaOH-Po contents under different irrigation modes. The NaOH-Po value was 61.16 mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e in the control group (C treatment), while this value was 56.48 mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e in alone aerated treatment (A), showing a 7.66% decrease compared to the control group (C). The CRI, CFM, and mixed inoculation (CFMRI) treatments reduced NaOH-Po by 22.15%, 20.74%, and 10.66%, respectively than control group (C). Considering the aerated irrigation treatments, ARI, AFM, and AFMRI treatments reduced NaOH-Po by 33.04%, 29.2%, and 15.79%, respectively, compared to alone aerated treatment (A). Combining RI, FM, or the RI\u0026thinsp;+\u0026thinsp;FM mixed inoculation with aerated drip irrigation further reduced NaOH-Po by 20.57%, 17.52%, and 12.96%, respectively, compared to the CRI, CFM, and CFMRI treatments. The combined use of ADI and AMF showed a significant synergistic effect, with the best results observed in the ADI\u0026thinsp;+\u0026thinsp;RI inoculant combination (ARI).\u003c/p\u003e\u003cp\u003eOverall, both ADI and AMF technologies reduced the total moderately available phosphorus contents in the soil. Compared to the control group values (145.19 mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), soil moderately available P content (137.95 mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) in aerated drip irrigation (A) was reduced by 5.25%. Inoculation with AMF also showed similar effects: for example, under the standard drip irrigation condition, RI, FM, and mixed inoculation reduced the moderately available P content by 14.61%, 12.18%, and 8.62%, respectively, as compared to the C treatment. Under oxygen-mycorrhiza synergy conditions, this effect was more pronounced, with ARI, AFM, and AFMRI treatments reduced the total contents of moderately available phosphorous by 22.93%, 20.04%, and 12.79%, respectively, compared to the C treatment.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec14\" class=\"Section3\"\u003e\u003ch2\u003e3.3.3 Insoluble phosphorus (Insoluble P)\u003c/h2\u003e\u003cp\u003eInsoluble forms of phosphorus (D-HCl-P, C-HCl-Pi/Po, and Residual-P) are typically not directly available to plants and can only be released as available phosphorus through long-term soil changes or microbial activity (Xu et al. 2020). The results regarding effects of different irrigation modes and mycorrhizal fungi inoculation on insoluble P fractions are given in Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eD-HCl-P is a phosphorus fraction extracted with diluted hydrochloric acid, primarily in the form of apatite and calcium-bound phosphorus. We found that, the D-HCl-P value was 252.57 mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e in control group (C treatment), while aerated drip irrigation treatment (A) showed a value of 237.45 mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, a 5.99% decrease compared to the control group (C). The CRI, CFM, and mixed inoculation (CFMRI) treatments reduced D-HCl-P by 11.96%, 12.83%, and 8.36%, respectively, compared to control group (C). When inoculation was done with RI, FM, or the RI\u0026thinsp;+\u0026thinsp;FM mixed inoculation under aerated drip irrigation, it was found that D-HCl-P was further reduced by 10.87%, 6.74%, and 6.03% than CRI, CFM, and CFMRI, respectively. In summary, both aerated drip irrigation and AMF inoculation led to a reduction in soil D-HCl-P (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;0.05), with no significant synergistic effect when combined.\u003c/p\u003e\u003cp\u003eC-HCl-Pi is stable inorganic phosphorus extracted with concentrated hydrochloric acid. There was no significant effect of aeration and fungi inoculation on C-HCl-Pi. Overall, it was observed that inoculation increased the C-HCl-Pi under both aerated and non-aerated condition. However, aerated drip irrigation has higher values for C-HCl-Pi than non-aerated irrigation mode (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;0.05). C-HCl-Po is stable organic phosphorus extracted with concentrated hydrochloric acid. We noticed that the aerated drip irrigation treatment (A) showed a 14.48% decrease in C-HCl-Po value compared to the control group (C). Inoculation with fungi also reduced the C-HCl-Po, such as CRI, CFM, and mixed inoculation (CFMRI) treatments have 19.15%, 13.55%, and 8.15% less C-HCl-Po compared to the control group (C) respectively. Besides, ARI, AFM, and AFMRI further reduced C-HCl-Po by 25.37%, 19.54%, and 13.64%, respectively as opposed to CRI, CFM, and CFMRI respectively. Briefly, synergistic effect of both aerated drip irrigation and AMF inoculation reduced the soil C-HCl-Po content (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05), with the highest reduction observed in ADI\u0026thinsp;+\u0026thinsp;RI inoculant combination (ARI).\u003c/p\u003e\u003cp\u003eResidual-P is soil residual phosphorus extracted by strong acid digestion. The Residual-P value in the control group (C treatment) was 104.49 mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e and was 101.28 mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e in aerated drip irrigation treatment (A), showing that aerated irrigation reduced Residual-P. The inoculation also notably minimized the Residual-P in soil, for example CRI, CFM, and mixed inoculation (CFMRI) treatments reduced Residual-P by 6.19%, 6.41%, and 10.78% respectively than control group (C). Furthermore, synergistic effect of ADI and AMF further decreased the Residual-P by 6.48%, 2.44%, and 3.52% than CRI, CFM, and CFMRI respectively (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05).\u003c/p\u003e\u003cp\u003eOverall, both ADI and AMF technologies slightly reduced the content of total insoluble phosphorus fractions in the soil. Compared to the control group contents (493.45 mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), soil insoluble P content in aerated drip irrigation (A) reduced by 3.55%. Inoculation with AMF showed similar decreasing effects, such as RI, FM, and mixed inoculation reduced the total insoluble P content by 5.18%, 6.74%, and 4.68%, respectively. Under oxygen-mycorrhiza synergy conditions, this effect in lowering the total insoluble phosphorus fraction was enhanced, with ARI, AFM, and AFMRI treatments reducing it by 11.94%, 9.63%, and 6.69% respectively compared to the control group. Further, ADI\u0026thinsp;+\u0026thinsp;RI inoculant (ARI) combination showed the best effect.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec15\" class=\"Section3\"\u003e\u003ch2\u003e3.3.4 Soil acid phosphatase activity\u003c/h2\u003e\u003cp\u003eThe results regarding soil acid phosphatase (Acid Phosphatase, ACP) activity measured in \u0026micro;mol p-nitrophenol\u0026middot;g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e dry soil\u0026middot;h\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e are given in Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e. The findings revealed that ACP value in the control group (C treatment) was 5.12 \u0026micro;mol\u0026middot;g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u0026middot;h\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e. While in aerated drip irrigation treatment (A), the ACP value was 7.2 \u0026micro;mol\u0026middot;g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u0026middot;h\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, which represented a 40.63% increase compared to the control group (C). The CRI, CFM, and mixed inoculation (CFMRI) treatments increased ACP activity by 54.81%, 25.98%, and 12.96%, respectively, compared to C treatment. Furthermore, the corresponding values for ACP activity under ARI, AFM, and AFMRI were high by 29.38%, 26.05%, and 35.22%, respectively, than CRI, CFM, and CFMRI inoculation treatments. Overall, combined use of ADI and AMF inoculation exhibited a synergistic effect, with the maximum increase observed in the ADI\u0026thinsp;+\u0026thinsp;RI inoculant combination (ARI).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003c/div\u003e\u003c/div\u003e\u003cdiv id=\"Sec16\" class=\"Section2\"\u003e\u003ch2\u003e3.4 Phosphorus accumulation in different plant organs, yield, and phosphorus use efficiency\u003c/h2\u003e\u003cp\u003eThe effects of ADI and AMF inoculation on phosphorus accumulation in the different organs of tomato plants are given in Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003e. Both ADI and AMF inoculation significantly affected phosphorus accumulation in the root, stem, leaf, and tomato fruits (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05). Overall, the phosphorus content in tomato organs followed the order of leaf\u0026thinsp;\u0026gt;\u0026thinsp;fruit\u0026thinsp;\u0026gt;\u0026thinsp;stem\u0026thinsp;\u0026gt;\u0026thinsp;root. The total phosphorus accumulation in aerated drip irrigation treatment (A) was 216.44 mg.pot\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, representing a 30.82% increase as compared to control group (C) value (165.45 mg\u0026middot;pot\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e). The CRI, CFM, and mixed inoculation (CFMRI) treatments increased total phosphorus accumulation by 22.94%, 15.09%, and 9.47%, respectively than the treatment without any inoculation and aeration (C). Besides, the corresponding values for total phosphorus accumulation under ARI, AFM, and AFMRI were further increased under aerated irrigation by 34.38%, 25.73%, and 21.37%, respectively as compared to CRI, CFM, and CFMRI. In summary, both aerated drip irrigation and AMF inoculation showed a significant synergistic effect which increased the total phosphorus accumulation in tomato plants, with the highest total phosphorus accumulation observed in the ADI\u0026thinsp;+\u0026thinsp;RI inoculant combination (ARI). Under the ARI treatment, the phosphorus accumulation in the stem, leaf, and fruit of tomatoes increased by 55.42%, 72.39%, and 80.66%, respectively, as opposed to the control group (C).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eThe effects of ADI and AMF inoculation on tomato yield (fresh weight) and mycorrhizal attributes are presented in Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e. We noticed that ADI significantly improved tomato yield (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05). The descending trend (from highest to lowest) for tomato yield was as follows: ARI\u0026thinsp;\u0026gt;\u0026thinsp;AFM\u0026thinsp;\u0026gt;\u0026thinsp;AFMRI\u0026thinsp;\u0026gt;\u0026thinsp;A\u0026thinsp;\u0026gt;\u0026thinsp;CRI\u0026thinsp;\u0026gt;\u0026thinsp;CFM\u0026thinsp;\u0026gt;\u0026thinsp;CFMRI\u0026thinsp;\u0026gt;\u0026thinsp;C. The aerated drip irrigation treatment (A) yielded 57.08 t\u0026middot;ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, which was 26.53% more yield as compared to the control treatment (C) that produced 45.12 t\u0026middot;ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e. Inoculation without aerated irrigation improved yield by 16.58%, 14.36%, and 10.83% in CRI, CFM, and CFMRI respectively than control (C). While ARI, AFM, and AFMRI treatments increased yield by 26%, 14.24%, and 10.76%, respectively, compared to the aerated treatment without inoculation (A). Overall, both aerated drip irrigation and AMF inoculation promoted an increase in tomato yield, with the highest yield observed in the ADI\u0026thinsp;+\u0026thinsp;RI inoculant combination (ARI).\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eTomato yield, mycorrhizal effects, and phosphorus use efficiency.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"6\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eTreatment\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eYield\u003c/em\u003e\u003c/p\u003e\u003cp\u003e(t\u0026middot;ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cem\u003eMGR\u003c/em\u003e\u003c/p\u003e\u003cp\u003e(%)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eMPR\u003c/em\u003e\u003c/p\u003e\u003cp\u003e(%)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u003cem\u003ePUE\u003c/em\u003e\u003c/p\u003e\u003cp\u003e(kg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cem\u003ePAE\u003c/em\u003e\u003c/p\u003e\u003cp\u003e(mg\u0026middot;g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e45.12\u0026thinsp;\u0026plusmn;\u0026thinsp;3.87d\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1038.06\u0026thinsp;\u0026plusmn;\u0026thinsp;26.16a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e28.91\u0026thinsp;\u0026plusmn;\u0026thinsp;2.86c\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCRI\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e52.6\u0026thinsp;\u0026plusmn;\u0026thinsp;2.08cd\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e15.19\u0026thinsp;\u0026plusmn;\u0026thinsp;3.02b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e18.52\u0026thinsp;\u0026plusmn;\u0026thinsp;3.41cd\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e985.16\u0026thinsp;\u0026plusmn;\u0026thinsp;73.04a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e34.9\u0026thinsp;\u0026plusmn;\u0026thinsp;5.28bc\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCFM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e51.6\u0026thinsp;\u0026plusmn;\u0026thinsp;2.3cd\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e14.05\u0026thinsp;\u0026plusmn;\u0026thinsp;3.74b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e12.92\u0026thinsp;\u0026plusmn;\u0026thinsp;4.12cd\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1030.46\u0026thinsp;\u0026plusmn;\u0026thinsp;40.89a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e29.89\u0026thinsp;\u0026plusmn;\u0026thinsp;4.49c\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCFMRI\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e50\u0026thinsp;\u0026plusmn;\u0026thinsp;2.28cd\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e9.91\u0026thinsp;\u0026plusmn;\u0026thinsp;4.46b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e7.93\u0026thinsp;\u0026plusmn;\u0026thinsp;4.03d\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1053.81\u0026thinsp;\u0026plusmn;\u0026thinsp;61.07a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e25.88\u0026thinsp;\u0026plusmn;\u0026thinsp;2.69c\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eA\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e57.08\u0026thinsp;\u0026plusmn;\u0026thinsp;5.42bc\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1004.19\u0026thinsp;\u0026plusmn;\u0026thinsp;68.71a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e35.01\u0026thinsp;\u0026plusmn;\u0026thinsp;3.1bc\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eARI\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e71.93\u0026thinsp;\u0026plusmn;\u0026thinsp;7.7a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e35.16\u0026thinsp;\u0026plusmn;\u0026thinsp;7.29a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e43.31\u0026thinsp;\u0026plusmn;\u0026thinsp;2.53a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e997.27\u0026thinsp;\u0026plusmn;\u0026thinsp;59.38a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e47.91\u0026thinsp;\u0026plusmn;\u0026thinsp;7.93a\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eAFM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e65.21\u0026thinsp;\u0026plusmn;\u0026thinsp;0.47ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e30.28\u0026thinsp;\u0026plusmn;\u0026thinsp;0.63a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e30.83\u0026thinsp;\u0026plusmn;\u0026thinsp;2.19b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1035.82\u0026thinsp;\u0026plusmn;\u0026thinsp;29.25a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e45.37\u0026thinsp;\u0026plusmn;\u0026thinsp;7.93ab\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eAFMRI\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e63.22\u0026thinsp;\u0026plusmn;\u0026thinsp;3.31ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e26.69\u0026thinsp;\u0026plusmn;\u0026thinsp;4.16a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e24.12\u0026thinsp;\u0026plusmn;\u0026thinsp;6.75bc\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1096.98\u0026thinsp;\u0026plusmn;\u0026thinsp;51.28a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e37.19\u0026thinsp;\u0026plusmn;\u0026thinsp;5.31abc\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eData are presented as mean\u0026thinsp;\u0026plusmn;\u0026thinsp;standard deviation. Values with the same lower case were not significant at \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05 among different treatments. C represents standard drip irrigation, and A represents aerated drip irrigation. RI refers to the treatment with \u003cem\u003eRhizophagus intraradices\u003c/em\u003e (RI) inoculation only; FM refers to the treatment with \u003cem\u003eFunneliformis mosseae\u003c/em\u003e (FM) inoculation only; FMRI refers to the mixed inoculation of FM and RI.\u003c/p\u003e\u003cp\u003eAdditionally, aeration significantly enhanced the mycorrhizal growth response (\u003cem\u003eMGR\u003c/em\u003e) and mycorrhizal phosphorus uptake response (\u003cem\u003eMPR\u003c/em\u003e) in studied tomatoes (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.01). The analysis showed that under standard drip irrigation conditions (C), the CRI treatment had the highest \u003cem\u003eMGR\u003c/em\u003e and \u003cem\u003eMPR\u003c/em\u003e at 15.19% and 18.52%, respectively, which were higher than the 14.05% and 12.92% in the CFM treatment, while the mixed inoculation (CFMRI) treatment had the lowest \u003cem\u003eMGR\u003c/em\u003e and \u003cem\u003eMPR\u003c/em\u003e at 9.91% and 7.93%, respectively. Same trend was found under ADI conditions, with the ARI treatment showing the highest \u003cem\u003eMGR\u003c/em\u003e and \u003cem\u003eMPR\u003c/em\u003e at 35.16% and 43.31%, respectively. Hence, RI inoculation showed high mycorrhizal effects under both irrigation conditions, indicating that RI inoculation contributed more to tomato dry matter accumulation and phosphorus uptake compared to FM, while aeration significantly enhanced this inoculation effect.\u003c/p\u003e\u003cp\u003eThe phosphorus use efficiency indicators (\u003cem\u003ePAE\u003c/em\u003e and \u003cem\u003ePUE\u003c/em\u003e) for tomatoes are given in Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e. The aerated drip irrigation treatment (A) had \u003cem\u003ePAE\u003c/em\u003e value of 35.01 mg\u0026middot;g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, showing 21.12% increase as compared to control treatment (C) which has PAE value of 28.91 mg\u0026middot;g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e. The CRI and CFM treatments increased \u003cem\u003ePAE\u003c/em\u003e by 20.75% and 3.39%, respectively, than control (C). The ARI, AFM, and AFMRI treatments increased \u003cem\u003ePAE\u003c/em\u003e by 36.86%, 29.59%, and 6.24%, respectively as opposed to their control treatment without any inoculation (A). Briefly, both aerated drip irrigation and AMF inoculation significantly increased \u003cem\u003ePAE\u003c/em\u003e, and their combined use demonstrated a significant synergistic effect (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05), with the best performance observed in the ADI\u0026thinsp;+\u0026thinsp;RI inoculant combination (ARI).\u003c/p\u003e\u003cp\u003eThere were no significant differences among various treatments regarding \u003cem\u003ePUE\u003c/em\u003e (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;0.05). The aerated drip irrigation treatment (A) had 3.26% decrease in PUE than control treatment (C). The CRI, CFM, and mixed inoculation (CFMRI) treatments affected \u003cem\u003ePUE\u003c/em\u003e in tomato by -5.1%, -0.73%, and 1.52%, respectively, compared to C treatment. The RI, FM, and RI\u0026thinsp;+\u0026thinsp;FM inoculation treatments under aerated drip irrigation increased \u003cem\u003ePUE\u003c/em\u003e by 1.23%, 0.52%, and 4.1%, respectively, compared to the CRI, CFM, and CRIFM. Overall, aerated drip irrigation slightly improved \u003cem\u003ePUE\u003c/em\u003e, while the effect of AMF inoculation on \u003cem\u003ePUE\u003c/em\u003e was inconsistent (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;0.05), because the best performance for \u003cem\u003ePUE\u003c/em\u003e was observed in the ADI\u0026thinsp;+\u0026thinsp;mixed inoculant combination (AFMRI) instead of ARI.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec17\" class=\"Section2\"\u003e\u003ch2\u003e3.5 Structural equation model of phosphorus accumulation and yield formation in greenhouse tomato\u003c/h2\u003e\u003cp\u003eA structural equation model (SEM) was constructed to analyze the synergistic effects of aerated drip irrigation (ADI) and arbuscular mycorrhizal fungi (AMF) inoculation on soil phosphorus transformation, root morphology development, plant phosphorus accumulation, and tomato yield regulation (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003e). The variance explanation (R\u003csup\u003e2\u003c/sup\u003e) for each endogenous variable was as follows: soil phosphatase activity (61.0%), soil inorganic phosphorus (44.4%), soil organic phosphorus (46.1%), root morphology development (44.9%), plant phosphorus accumulation (68.4%), and yield (86.1%). These results indicate that the model has good explanatory power and integrates the key processes affecting tomato yield formation. The path analysis of the structural equation model revealed the following:\u003c/p\u003e\u003cp\u003e(\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e) \u003cb\u003eDrivers of soil phosphatase activity\u003c/b\u003e: Dissolved oxygen in irrigation water (path coefficient\u0026thinsp;=\u0026thinsp;1.219, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001) and AM fungi (path coefficient\u0026thinsp;=\u0026thinsp;0.517, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001) both significantly and positively influenced soil phosphatase activity, indicating that both factors synergistically promoted the activation of phosphatase.\u003c/p\u003e\u003cp\u003e(\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e) \u003cb\u003eRegulation of phosphorus fractions by phosphatase\u003c/b\u003e: We found that soil phosphatase activity significantly and positively drove soil inorganic phosphorus (path coefficient\u0026thinsp;=\u0026thinsp;0.666, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), while negatively regulated soil organic phosphorus (path coefficient = -0.679, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001). This reflects that phosphatase accelerates organic phosphorus mineralization and enriches inorganic phosphorus.\u003c/p\u003e\u003cp\u003e(\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e) \u003cb\u003eResponse of root morphology development\u003c/b\u003e: Dissolved oxygen in irrigation water (path coefficient\u0026thinsp;=\u0026thinsp;1.030, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001) and AM fungi (path coefficient\u0026thinsp;=\u0026thinsp;0.452, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.01) inoculation both significantly and positively shaped root morphology development, indicating that ADI may improve root morphology development by alleviating root hypoxic stress, while AM fungi might optimize root morphology through hyphal extension and signaling regulation.\u003c/p\u003e\u003cp\u003e(\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e) \u003cb\u003eFactors influencing plant phosphorus accumulation\u003c/b\u003e: Soil organic phosphorus contents were negatively associated with plant phosphorus accumulation (path coefficient = -0.451, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05), while root morphology development significantly promoted phosphorus accumulation (path coefficient\u0026thinsp;=\u0026thinsp;0.365, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05). Besides, soil inorganic phosphorus benefited plant phosphorus accumulation, but this effect was not significant (path coefficient\u0026thinsp;=\u0026thinsp;0.103, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;0.05). These three factors together explained 68.4% (R\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.684) of the variation in plant phosphorus accumulation. Briefly, oxygen-mycorrhiza synergy might promote phosphorus accumulation by enhancing soil phosphatase activity, accelerating organic phosphorus mineralization, and regulating inorganic phosphorus supply, as well as optimizing root morphology to directly promote phosphorus absorption.\u003c/p\u003e\u003cp\u003e(\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e) \u003cb\u003eDominant pathways in yield formation\u003c/b\u003e: Plant phosphorus accumulation significantly and positively determined yield (path coefficient\u0026thinsp;=\u0026thinsp;0.911, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), explaining 86.1% (R\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.861) of the variation in yield. Root morphology development had no significant direct effect on yield (path coefficient\u0026thinsp;=\u0026thinsp;0.023, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;0.05), indicating that yield improvement mainly relies on the indirect effects of efficient phosphorus accumulation.\u003c/p\u003e\u003cp\u003eThe model reliability and validity tests showed that most of the latent variables had Cronbach's α\u0026thinsp;\u0026gt;\u0026thinsp;0.8, while all variables have average variance extracted (AVE)\u0026thinsp;\u0026gt;\u0026thinsp;0.5, and composite reliability (CR)\u0026thinsp;\u0026gt;\u0026thinsp;0.7, confirming that measurement model has good internal consistency and convergent validity. In conclusion, ADI and AMF synergistically improve soil phosphatase activity, promote organic phosphorus mineralization and inorganic phosphorus supply, which leads to optimize root morphology, ultimately increasing plant phosphorus accumulation and enhancing yield. Further, soil phosphatase plays a central role in phosphorus transformation, and plant phosphorus accumulation is a key mediator of yield formation.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003c/div\u003e"},{"header":"4. Discussion","content":"\u003cp\u003eThis study systematically explored the synergistic effects of aerated drip irrigation (ADI) and AMF inoculation on soil phosphorus transformation, root morphology, phosphorus accumulation, and yield. The results showed that the combination of ADI and \u003cem\u003eRhizophagus intraradices\u003c/em\u003e (RI) inoculation (ARI treatment) exhibited significant synergistic effects in enhancing AMF colonization, improving root morphology, promoting soil phosphorus transformation, and increasing phosphorus uptake and yield in tomatoes. This study therefore introduces the concept of \"Oxygen-Mycorrhiza Synergy\".\u003c/p\u003e\u003cdiv id=\"Sec19\" class=\"Section2\"\u003e\u003ch2\u003e4.1 Oxygen-mycorrhiza synergy on soil phosphorus fractions and availability\u003c/h2\u003e\u003cp\u003eThe availability of soil phosphorus directly impacts plant phosphorus uptake and growth. The transformation of phosphorus fractions is a key process in enhancing phosphorus availability. Based on the Hedley phosphorus fractionation system, this study found that the effects of ADI and AMF inoculation on soil phosphorus fractions followed the gradient pattern of \"easily available phosphorus\u0026thinsp;\u0026gt;\u0026thinsp;moderately available phosphorus\u0026thinsp;\u0026gt;\u0026thinsp;insoluble phosphorus\". The AMF inoculation increased easily available phosphorus contents in soil (Resin-P and NaHCO\u003csub\u003e3\u003c/sub\u003e-Pi), while decreasing the organic phosphorus components (such as NaHCO\u003csub\u003e3\u003c/sub\u003e-Po and NaOH-Po). However, the short-term impact of AMF on stable phosphorus fractions in the soil (HCl-P, Residue-P) was minimal. Phosphatases play a critical role in the microbial-mediated conversion of insoluble phosphorus (Lidbury et al. 2022), and this transformation indicates that AMF promoted organic phosphorus mineralization through the secretion of acid phosphatase, thereby enhancing phosphorus availability in the soil. This phenomenon aligns with the results observed in our study, where organic phosphorus decreased, and available phosphorus increased, as AMF hyphae exhibit higher acid phosphatase activity in the presence of organic phosphorus (Huang et al. 2023).\u003c/p\u003e\u003cp\u003eThe oxygen-mycorrhiza synergy amplified this transformation effect and significantly altered the distribution of soil phosphorus. Under the ARI treatment, soil acid phosphatase activity was 29.38% higher than that in the CRI treatment, while easily available phosphorus increased by 5.83%. Besides, moderately available phosphorus and insoluble phosphorus decreased by 9.74% and 7.13% respectively. The possible reasons for these changes could be: (\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e) ADI may have selectively enriched microbial groups related to phosphorus transformation (such as Proteobacteria and Acidobacteria) by altering soil redox conditions, further optimizing phosphorus cycling pathways (Gross et al. 2020; Kim et al. 2021); (\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e) Hypoxia could reduce the abundance of phosphate-dissolving bacteria such as Pseudomonas and Bacillus (Cui et al. 2022), and these bacteria usually dissolve insoluble phosphate by secreting organic acids and enzymes (Ouyang et al. 2021). As ADI increased dissolved oxygen in the irrigation water and soil porosity, thereby improving soil gas exchange, which likely enhanced the phosphorus-solubilizing bacteria and AMF metabolic activity. The organic acids (such as citric acid and oxalic acid) secreted by phosphorus-solubilizing bacteria lower soil pH, chelate metal ions like Fe\u003csup\u003e3+\u003c/sup\u003e and Al\u003csup\u003e3+\u003c/sup\u003e, and dissolve certain phosphates (e.g., Fe-P, Al-P) (Pang et al. 2024; Aliyat et al. 2022). Meanwhile, AMF upregulates the expression of genes related to host plant acid phosphatase secretion, which increases soil acid phosphatase activity and accelerates organic phosphorus mineralization (Nopphakat et al. 2022; Pang et al. 2024). As a result, oxygen-mycorrhiza synergy enhanced the availability of phosphorus to tomatoes. This finding has significant implications for optimizing soil phosphorus management and improving phosphorus fertilizer use efficiency.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec20\" class=\"Section2\"\u003e\u003ch2\u003e4.2 Effects of oxygen-mycorrhiza synergy on tomato-mycorrhizal symbiosis and phosphorus uptake\u003c/h2\u003e\u003cp\u003eMycorrhizal symbiosis is an important pathway for plants to acquire mineral nutrients, especially under low phosphorus availability in soil. The colonization rate is a key indicator of the effectiveness of mycorrhizal symbiosis. In our study, ADI significantly increased tomato mycorrhizal colonization, particularly after inoculation with \u003cem\u003eRhizophagus intraradices\u003c/em\u003e (RI), which showed a 32.66% increase in colonization rate compared to conventional drip irrigation without aeration. This may be closely related to the improvement of rhizosphere oxygen conditions by ADI, which facilitated spore germination and hyphal invasion. Suitable soil oxygen concentrations are crucial for AMF growth and activity, as AMF are aerobic microorganisms whose metabolic activities depend on oxygen supply (Zhang et al. 2014). Xu et al. (2021) found that aeration significantly increased dissolved oxygen concentrations and AMF colonization in wetland ecosystems, promoting plant growth. In current study, intermittent aeration (4 hours\u0026middot;day\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) might stabilized the aerobic environment, facilitating AMF colonization of roots. Furthermore, ADI improved soil aeration, promoted root respiration, and increased the secretion of carbohydrates (such as sucrose and glucose) from roots (Zheng et al. 2024), providing more carbon sources for AMF, thereby accelerating the formation of hyphal networks. Moreover, aeration may enhance the plant's ability to recognize and colonize AMF by modulating plant hormone balance (such as auxins and abscisic acid) (Chareesri et al. 2020). Notably, RI inoculation showed higher colonization efficiency than \u003cem\u003eFunneliformis mosseae\u003c/em\u003e (FM) and the mixed inoculation (FM\u0026thinsp;+\u0026thinsp;RI), suggesting potential competition effects between different AMF strains (Berruti et al. 2016).\u003c/p\u003e\u003cp\u003eOxygen-mycorrhiza synergy also optimized tomato root morphology. The AMF inoculation remarkably improved plant nutrient uptake by changing root branching patterns and increasing the proportion of fine roots (Chandrasekaran 2022). In this study, soil aeration significantly influenced tomato root length, root surface area, and root tip number (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05). Specifically, under the ARI treatment, the total root length, root surface area, and root branching increased by 23.65%, 19.96%, and 23.67%, respectively, compared to the CRI treatment. In contrast, the average root diameter ranged from 1.10 to 1.27 mm, with no significant differences observed between treatments. The effect of aeration on root diameter was not significant (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;0.05), which may be attributed to the fact that although ADI increased root volume, it also promoted root growth and elongation (as indicated by increased root tip number and total root length), thus exerting no significant impact on root diameter. However, it also suggests that tomato roots have a larger specific surface area, more branching, and finer root hairs, which facilitate better interaction with soil nutrients (Nascimento et al. 2021). Furthermore, ADI improved root development by enhancing soil oxygen supply, as oxygen is essential for root growth and metabolic activities (Roosta 2024; Wang et al. 2023b). The synergistic effects of ADI and AMF to amplify the improvement in root morphology might be attributed to ADI alleviating soil compaction and hypoxic stress, enhancing root vitality and respiration efficiency. The improved root structure provided more colonization sites for AMF, and the enhanced nutrient absorption capacity of AMF promoted denser lateral root development, thereby increasing surface area and forming a more efficient material exchange interface. The core mechanism of oxygen-mycorrhiza synergy lies in optimizing the rhizosphere microenvironment, activating fungal functional expression, and reshaping root growth patterns, ultimately affecting root absorption and supporting functions (Zeng et al. 2025; Wang et al. 2023a).\u003c/p\u003e\u003cp\u003eBoth aerated drip irrigation and AMF inoculation significantly increased tomato phosphorus accumulation, consistent with previous studies (Bowles et al. 2016). Under oxygen-mycorrhiza synergy, higher phosphorus availability and improved root morphology led to significant increases in phosphorus accumulation in tomato stems, leaves, and fruits. In this study, the ARI treatment showed a 34.38% higher phosphorus accumulation in fruit compared to CRI. Furthermore, the ARI treatment achieved the highest yield (71.93 t\u0026middot;ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), a 36.75% increase compared to CRI. Similar findings were also reported by Leventis et al. (2021) in controlled experiments with tomato inoculation (Leventis et al. 2021). The reasons behind the increase in tomato productivity under combined use of ADI and AMF could be: AMF may break through the phosphorus-depleted zones in the rhizosphere with its hyphal networks, transferring phosphorus to the host plant and expanding the root nutrient absorption range (Wen et al. 2020); while aerated drip irrigation improves the soil microenvironment, enhancing AMF activity and colonization, thus interactively promoting phosphorus uptake by plants. It is important to note that different AMF inoculation modes contributed differently to tomato phosphorus accumulation and yield. The mycorrhizal effect of RI inoculation was significantly high than that of FM inoculant and mixed inoculation, which may be due to the stronger adaptability of the RI strain to the tested soil and oxygen environment. In mixed inoculation, the growth-promoting effect of FM and RI may be weakened or even antagonistic due to resource competition, niche overlap, or metabolic interference (Zhang et al. 2024), although still higher than the control group without inoculation. Additionally, symbiosis is often fine-tuned based on plant needs and surrounding conditions, typically through plant hormone signaling. Lidoy et al. (2023) compared the colonization ability of \u003cem\u003eFunneliformis mosseae\u003c/em\u003e and \u003cem\u003eRhizophagus irregularis\u003c/em\u003e under hormone and salt stress conditions, concluding that colonization levels depend on the fungal genotype and stress conditions. In this study, soil aeration may have influenced the phosphorus signaling pathways of tomato plants by modulating soil oxygen stress, preferentially activating gene expression for symbiosis with RI, thereby enhancing the functional advantage of the specific fungal strain. This was reflected in the best symbiotic effect observed with the RI inoculant in tomato plants.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec21\" class=\"Section2\"\u003e\u003ch2\u003e4.3 Mechanism involved in oxygen-mycorrhiza synergy to increase tomato yield and phosphorus acquisition\u003c/h2\u003e\u003cp\u003eThe SEM revealed the potential mechanisms by which ADI and AMF synergistically enhance yield formation. The model showed that soil phosphatase activity is a key factor in phosphorus transformation and significantly positively influenced the soil inorganic phosphorus contents (path coefficient\u0026thinsp;=\u0026thinsp;0.666). This finding is consistent with research suggesting that AMF increase phosphorus availability by modulating soil enzyme activity (Sheteiwy et al. 2021), and we notably found that oxygen-mycorrhiza synergy mainly promotes the conversion of organic phosphorus to inorganic phosphorus. In this study, oxygen-mycorrhiza synergy optimized phosphorus supply by enhancing phosphatase activity and improving the structure of soil phosphorus fractions. Simultaneously, it promoted tomato root development (path coefficient\u0026thinsp;=\u0026thinsp;1.030) and enhanced the plant phosphorus uptake capacity (path coefficient\u0026thinsp;=\u0026thinsp;0.365). These results indicate that the combined application of aerated drip irrigation and AMF not only promotes effective phosphorus absorption by enhancing soil phosphatase activity and inorganic phosphorus supply, but also further improved phosphorus accumulation and yield in tomatoes by optimizing root morphology. Moreover, the model also revealed the significant positive impact of phosphorus accumulation on tomato yield (path coefficient\u0026thinsp;=\u0026thinsp;0.911), emphasizing the crucial role of phosphorus uptake in yield formation. Finally, our findings underscore the importance of integrating soil biological processes and plant physiological responses in optimizing nutrient management and crop production.\u003c/p\u003e\u003c/div\u003e"},{"header":"5. Conclusion","content":"\u003cp\u003eThis study emphasized the critical role of oxygen-mycorrhiza synergy in regulating availability of soil phosphorous via complex interactions involving mineralization, colonization, and enzymatic activities. We found that aeration can optimize the symbiotic relationship between tomato roots and AMF, thereby significantly increasing mycorrhizal colonization rate. The oxygen-mycorrhiza synergy notably improved acid phosphatase activity, promoted the conversion of soil active organic phosphorus to active inorganic phosphorus, and facilitates the transformation of insoluble phosphorus to available phosphorus. Simultaneously, both ADI and AMF optimized tomato root morphology, jointly promoting phosphorus accumulation in plants and ultimately increasing yield. In summary, the combined use of ADI and AMF has a significant synergistic effect in promoting phosphorus uptake in tomatoes and improving the phosphorus availability in soils. Future studies should further explore the specific pathways and molecular mechanisms through which oxygen-mycorrhiza synergy influence soil phosphorus availability, integrating root exudates and microbial community data within the \"soil oxygen environment-microorganisms-plant physiology\" framework.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAvailability of data and materials:\u003c/strong\u003e The data that supports the findings of this study are available from the corresponding author upon reasonable request.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor contributions:\u003c/strong\u003e \u003cstrong\u003eYingji Lian\u003c/strong\u003e: Investigation, Data curation, Writing \u0026ndash; original draft, Visualization; \u003cstrong\u003eHongjun Lei\u003c/strong\u003e: Conceptualization, Resources, Writing \u0026ndash; review \u0026amp; editing, Supervision, Project administration, Funding acquisition; \u003cstrong\u003eHongwei Pan\u003c/strong\u003e: Writing \u0026ndash; review \u0026amp; editing, Resources; \u003cstrong\u003eMuhammad Zain\u003c/strong\u003e: Language polishing, Review \u0026amp; editing; \u003cstrong\u003eXin Liu\u003c/strong\u003e: Software support , Visualization; \u003cstrong\u003eShaobo Wang:\u003c/strong\u003e Writing \u0026ndash; review \u0026amp; editing; \u003cstrong\u003eYong Liu\u003c/strong\u003e: Experimental instruction, Supervision; \u003cstrong\u003eShihui Zhang\u003c/strong\u003e: Data analysis, Investigation.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e: This work was supported by the National Natural Science Foundation of China (52079052) and the PhD Innovation Fund of North China University of Water Resources and Electric Power (BCJJ202407).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConflicts of interest:\u003c/strong\u003e The authors declare that they have no conflict of interest.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAliyat F, Maldani M, Guilli M, et al. 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Agricultural water management 274: 107925. https://doi.org/10.1016/j.agwat.2022.107925\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":true,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Aerated drip irrigation, Arbuscular mycorrhizal fungi, Mycorrhizal colonization rate, Phosphorus transformation, Phosphorus accumulation","lastPublishedDoi":"10.21203/rs.3.rs-7489363/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7489363/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003ch2\u003eBackground and Aims:\u003c/h2\u003e\u003cp\u003eLimited soil phosphorus (P) availability impairs plant growth in protected cultivation systems. Although arbuscular mycorrhizal fungi (AMF) can enhance P uptake, the combined effects of AMF and aerated drip irrigation (ADI) on soil P fractions and tomato P accumulation remain unclear.\u003c/p\u003e\u003ch2\u003eMethods\u003c/h2\u003e\u003cp\u003eWe conducted a greenhouse experiment with two irrigation regimes\u0026mdash;standard drip (DO\u0026thinsp;=\u0026thinsp;6 mg\u0026middot;L⁻\u0026sup1;) and ADI (DO\u0026thinsp;=\u0026thinsp;15 mg\u0026middot;L⁻\u0026sup1;)\u0026mdash;and four AMF treatments: control, \u003cem\u003eFunneliformis mosseae\u003c/em\u003e (FM), \u003cem\u003eRhizophagus intraradices\u003c/em\u003e (RI), and FM\u0026thinsp;+\u0026thinsp;RI mixed inoculation. We assessed AMF colonization, soil P fractions, tomato P accumulation, yield, and employed structural equation modeling (SEM) to elucidate mechanisms.\u003c/p\u003e\u003ch2\u003eResults\u003c/h2\u003e\u003cp\u003eADI combined with RI (ARI) significantly enhanced AMF colonization (+\u0026thinsp;32.66%, P\u0026thinsp;\u0026lt;\u0026thinsp;0.05), increased the readily available Resin-P fraction by 32.09%, promoted organic-to-inorganic P conversion (NaHCO\u003csub\u003e3\u003c/sub\u003e-Pi\u0026thinsp;+\u0026thinsp;15.87%, NaHCO\u003csub\u003e3\u003c/sub\u003e-Po \u0026minus;\u0026thinsp;23.64%, NaOH-Po \u0026minus;\u0026thinsp;20.57%), and resulted in 34.4% greater plant P accumulation and 36.8% higher yield compared to RI under standard irrigation. SEM revealed two key mechanisms: increased acid phosphatase activity driving organic P mineralization and optimized root morphology enhancing P uptake.\u003c/p\u003e\u003ch2\u003eConclusion\u003c/h2\u003e\u003cp\u003eOur findings support that ADI and AMF synergistically improve P availability, uptake, and tomato productivity by integrating soil biochemical transformation and enhanced root architecture\u0026mdash;offering a promising strategy for sustainable phosphorus management in greenhouse production.\u003c/p\u003e","manuscriptTitle":"Oxygen-mycorrhiza synergy drives phosphorus transformation in soil and its accumulation in greenhouse-grown tomato.","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-09-08 17:49:16","doi":"10.21203/rs.3.rs-7489363/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"2e899e46-8db9-4ec9-af75-b25f3f028d0d","owner":[],"postedDate":"September 8th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2026-01-15T11:10:01+00:00","versionOfRecord":[],"versionCreatedAt":"2025-09-08 17:49:16","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-7489363","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7489363","identity":"rs-7489363","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}