Reductive soil disinfestation promoted vegetable N uptake by regulating soil gross N transformations and improving the quality of degraded vegetable soils

Reductive soil disinfestation promoted vegetable N uptake by regulating soil gross N transformations and improving the quality of degraded vegetable soils | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Reductive soil disinfestation promoted vegetable N uptake by regulating soil gross N transformations and improving the quality of degraded vegetable soils jinbo zhang, Xiaoqian Dan, Mengqiu He, Shending Chen, Xiaoxiang He, and 4 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-1805779/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 6 You are reading this latest preprint version Abstract Aims Reductive soil disinfestation (RSD) has been widely applied to improve soil degradation, thereby enhancing vegetable N uptake and subsequently productivity. However, the effect of RSD on interactions between vegetable N uptake and soil gross N transformation remain unclear. Methods Two degraded vegetable soils were treated with RSD. Untreated soils served as control (CK). To quantify the effects of RSD on N cycling in vegetable-soil systems, 15 N tracing pot experiments were conducted. Results Vegetable NH 4 + and NO 3 − uptake rates were 1.0–6.5 times higher in RSD treatments than CK. Soil gross N mineralization rates ( M ) in RSD ranged from 0.83 to 13.00 mg N kg − 1 d − 1 , and were significantly higher than in CK (0.21 to 8.71 mg N kg − 1 d − 1 ). Autotropic nitrification rates ( O NH4 ) increased by 1.7–4.2 times after RSD. NH 4 + immobilization rates ( I NH4 ) were significantly inhibited by RSD in the presence of vegetables. These induced decreasing ( O NH4 + I NH4 )/ M , increasing NH 4 + retention times and production rates of soil NO 3 − after RSD treatment. Thus, RSD promoted N supply to vegetables and subsequently N uptake by vegetables on degraded soils. In addition, RSD improved the quality on degraded soils (i.e. increasing soil pH, and decreasing soil EC and NO 3 − contents and pathogen abundances), which could also be an important factor promoting vegetable N uptake. Conclusion Thus, RSD can promote vegetable N uptake by regulating soil gross N transformations and improving the quality of degraded vegetable soils. However, N fertilizer management still needs attention because of stimulated NO 3 − production rates after RSD, which may lead to more rapid NO 3 − leaching or gaseous N losses. Reductive soil disinfestation vegetable N uptake Gross N transformation rates degraded vegetable soils 15N tracing Figures Figure 1 Figure 2 Figure 3 Introduction Due to the high demand in the markets, the greenhouse vegetable production (GVP) is increasing rapidly worldwide. By 2018, the global area of GVP was approximately 5.6 million ha (Cuesta 2019 ; Fei et al. 2018 ). For high yield and economic profit, agricultural practices, such as intensive crop rotations (Jia et al. 2012 ) and high rates of N fertilizer application (often exceeding 1000 kg N ha-1, Ju et al. 2006 ; Shi et al. 2009 ), are widespread, especially in GVP. However, low N absorption capacity arising from the sparse and shallow root systems and short growth cycle of vegetables induce low N use efficiency (NUE) in GVP, which is generally < 20% (Ren et al. 2010 ; Ti et al. 2015 ). Consequently, long-term GVP lead to soil degradation, e.g. accumulation of NO 3 − in the soil, acidification, salinization and severe soil-borne diseases (Guo et al. 2010 ; Hu et al. 2017 ; Huang et al. 2016 ). This deterioration in the quality, safety and yield of vegetables led to environment problems and jeopardized the sustainable development of GVP (Albornoz 2016 ; Hu et al. 2017 ). Reductive soil disinfestation (RSD), as an environmentally friendly method developed in Japan and Netherlands in 2000 (Blok et al. 2000 ; Shinmura 2000 ), has been widely promoted to improve the quality of degraded vegetable soils (Butler et al. 2012b ; Huang et al. 2016 ; Li et al. 2019 ). Briefly, in the RSD method, easily decomposable organic material (e.g. molasses, alfalfa, maize straw) is homogeneously added into the soil. The soil is then irrigated until saturation, covered with plastic film and cultivated at a temperature of 30 to 35°C for 2−4 weeks. This treatment creates strong reductive conditions and high temperatures to effectively improve the physical, chemical and biological properties of degraded vegetable soils (Butler et al. 2012b ; Huang et al. 2016 ; Liu et al. 2018 ). For instance, the addition of organic matter can improve soil structure (such as enhancements of soil holding water and ions exchange), in favor of vegetable growth (Akhtar and Malik 2000; Oka 2010 ). Combining strong reductive condition with easily decomposable organic matter amendment can simulate the growth of soil denitrification microorganisms (e.g. nirK , nirS and nosZ ) (Huang et al. 2016 ; Huang et al. 2019 ; Zhao et al. 2020 ), which can promote the denitrification of NO 3 − and further rapidly remove soil accumulated NO 3 − . In addition, RSD can decrease soil electrical conductivity (EC), enhance soil pH and NH 4 + content, and efficiently suppress soil-borne plant pathogens and plant-parasitic nematodes (Butler et al. 2012a ; Huang et al. 2016 ; Liu et al. 2018 ; Zhao et al. 2018 ). Thus, degraded vegetable soils treated with RSD can provide a more favorable growing environment for vegetables and improve their N uptake, which promotes NUE and vegetable yield and reduces the environmental risk of N loss. In addition, the interactions between soil gross N transformations could affect plant N uptake, as they determine the amount and form of available N in the soil (Zhang et al. 2018 ). It is widely reported that RSD significantly alters soil properties (e.g. soil pH, total N, organic carbon (C), NH 4 + , NO 3 − , AOB, AOA, bacteria and fungi) (Huang et al. 2016 ; Zhu et al. 2014 ), which were key factors regulating soil gross N transformations. For instance, soil organic substrates (such total N and C) were vital energy sources for microorganisms and could drive microbial-mediated soil N cycling (e.g. soil gross N mineralization and heterotrophic nitrification) (Elrys et al. 2021a ; Elrys et al. 2021b ; Zhang et al. 2014 ; Zhang et al. 2019 ). Increasing soil pH can stimulate autotrophic nitrification by enhancing the release of NH 3 and the activities of ammonia-oxidizing microorganisms (e.g. AOA and AOB) (Li et al. 2018 ; Norton and Stark 2011 ; Stempfhuber et al. 2015 ). These were confirmed by Zhu et al. ( 2014 ) where RSD significantly enhanced heterotrophic nitrification and autotrophic nitrification by increasing soil pH, total N and organic carbon. In addition, the increase of soil microbial activity resulted in higher soil microbial immobilization rates after RSD. Although Zhu et al. ( 2014 ) concluded that RSD altered degraded vegetable soils gross N transformation rates. However, their results lacked the influence of vegetables N uptake, leading to an incomplete understanding of soil N transformations after RSD. Soil-plant feedbacks influence soil gross N transformations through transfer of organic matter into the soil in the form of litter or root exudates, which reduces the substrates of N transformations and affects the activity of microorganisms involved in N transformations (Dan et al. 2022 ; He et al. 2021 ; Nacry et al. 2013 ; Zhao et al. 2022 ). At present, the effects of RSD on the interactions between vegetable N uptake and soil gross N transformation rates are still unclear. A series of 15 N tracing pot experiments were carried out with vegetables to quantify the effects of RSD treatment on two degraded soils in terms of soil gross N transformation rates and plant N uptake rates using a 15 N tracing approach ( Ntrace plant ). Our study was carried out to test the following hypotheses: 1) soil gross N mineralization and autotrophic nitrification can be stimulated by RSD, considering that soil total N, pH and activities of ammonia-oxidizing microorganisms increased after RSD; 2) RSD can promote vegetable N uptake through effectively improving the quality of degraded vegetable soil. Material And Methods Soil samples Degraded greenhouse soils that had been under continuous cropping with tomato (Lycopersicon esculentum L.) (DST) and cucumber ( Cucumis sativus L.) (DSC) were sampled from greenhouses in Nanjing, Jiangsu Province, China. Soils were classified as udalfs (Alfisols) based to the soil taxonomy of the United States of America. On DST, tomato and strawberry had been grown continuously for more than ten years, leading to problems with continuous cultivation of tomatoes and strawberries. On DSC, more than 20 crops of continuous monoculture cultivation which resulted in an obstacle to continuous cucumber cultivation. Both two soils were collected after vegetable harvest. At each greenhouse, 4 plots with 1 m ×1 m were randomly selected, and surface soil (0-20 cm) was sampled and then mixed in equal amounts, subsequently sieved (2 mm) and roots were removed. Each soil was divided into two parts, one was treated by the reductive soil disinfestation (RSD) method, the other was store at 4 °C and served as a control (CK). Therefore, four soil samples, i.e. DST without and with RSD (named as DST-CK and DST-RSD, respectively) and DSC without and with RSD (named as DSC-CK and DSC-RSD, respectively), were used for the experiments. For RSD method, each soil was incorporated with alfalfa meal at a ratio of 1/50 (dry soil/alfalfa meal, w / w ), irrigated to 100% water hold capacity (WHC) and then evenly mixed. Treated soils were packed in zip-lock bags, sealed, placed in an incubator with 30 °C and cultivated for 21 days. Then, soils were dried to approximate 20% WHC, and passed through a 2 mm sieve. Each soil sample (whether RSD or not) was divided into 3 subsamples to determine the soil chemical (Table1), microbial properties (Table2) and conduct 15 N-tracing studies, respectively. 15 N-tracing experiments Eight treatments were setup, based on the four soil samples (e.g. DST-CK, DST-RSD, DSC-CK and DSC-RSD ) with and without plants, i.e. 1) only DST-CK soil (DST-CC), 2) DST-CK soil with tomato (DST-CP), 3) only DST-RSD soil (DST-RC), 4) DST-RSD soil with tomato (DST-RP), 5) only DSC-CK soil (DSC-CC), 6) DSC-CK soil with cucumber (DSC-CP), 7) only DSC-RSD soil (DSC-RC), and 8) DSC-RSD soil with cucumber (DSC-RP). Tomato and cucumber seeds were sterilized using 5% alcohol for 5 min, and washed 4 times using deionized water. The seeds were germinated for 2-3 days on petri dishes (with a moistened gauze on the bottom) placing in a 30 ° C-incubator. After germination the seedlings with uniform growth rate were selected and transplanted to the seeding plug tray. When its first true leaf had been fully developed, seedlings of uniform growth were transplanted into 360-mL pots containing 0.2 kg soil (oven-dry basis) after the nutritional material on roots was largely removed. Meanwhile, the treatments without planting were also set up. Each pot received nutrient solutions containing N, K and P with 50, 30 and 20 mg kg -1 soil, respectively, and trace elements, and was placed in a greenhouse for cultivation. The soil water content maintained at approximately 60% WHC by daily water additions. After 26 and 18 days of transplanting tomato and cucumber seedlings, respectively, a 15 N tracing experiment was carried out including all treatments. Two 15 N-labeled groups were established for each treatment, where either NH 4 + was 15 N labelled ( 15 NH 4 NO 3 , 10.20 atom% for all treatments) or where NO 3 – was 15 N labelled (NH 4 15 NO 3 , 10.12 atom% for DST-CC, DST-CP, DST-RC, DST-RP, DSC-RC, DSC-RP treatments, and 99.23 atom% for DSC-CC, DSC-CP treatments). 2-mL each 15 N solution was uniformly applied to each pot using a 4-needle injection technique at a rate of 30 mg NH 4 + -N kg -1 soil and 30 mg NO 3 – -N kg -1 soil for DST, and 50 mg NH 4 + -N kg -1 soil and 50 mg NO 3 – -N kg -1 soil (He et al. 2022). At 0.5, 24, 48 and 72 h after 15 N labelling, both soil and plants were collected. 1 M potassium chloride (KCl) solution (soil/solution with 1/5) was used to extract NH 4 + and NO 3 – in soil for determination of concentrations and 15 N abundances of NH 4 + and NO 3 – . Other soil properties, i.e. Soil pH, DOC (dissolved organic carbon), the bacteria, fungi, AOA (ammonia-oxidizing archaea) and AOB (ammonia-oxidizing bacteria) abundances, were also measured. Residual inorganic N on plant surfaces were thoroughly rinsed with 1 M KCl solution and then deionized water. After that, vegetables (including root) were de-enzymed in ovens with 105 °C for 0.5 h, dried in ovens with 80 o C to a constant weight. Dried vegetables were weighed to obtain their biomass, and then grinded and sieved (0.15 mm) to analyze their N concentrations and 15 N abundances. Analytical methods Soil pH (soil/water, 2.5:1) and EC (electrical conductivity) (soil/water, 5:1) were measured using DMP-2mV/pH detector (Quark Ltd., Nanjing, China) and conductivity detector (Kang Yi Crops., Nanjing, China), respectively. Soil organic carbon (SOC) and total N (TN) were analyzed by wet digestion with H 2 SO 4 -K 2 Cr 2 O 7 and semi-micro Kjeldahl digestion with Se, CuSO 4 , and K 2 SO 4 as catalysts, respectively. Soil dissolved organic C (DOC) content was determined using the Analyzer Multi N/C (Analytic Jena, Jena, Germany). NaOH fusion and Mo-Sb colorimetry, NaOH fusion and flame photometer, NaHCO 3 digestion and Mo-Sb colorimetry, as well as CH 3 COONH 4 digestion and flame photometer were used to analyze the total soil phosphorus (TP), total potassium (TK), available phosphorus (AP) and available potassium (AK), respectively, according to Bao (2000). Soils were extracted with 1 M KCl solution (soil to solution ratio, 1:5), the automated continuous flow wet chemistry analyzer (SA1000, Skalar, Netherlands) was used to analyze the concentrations of NH 4 + and NO 3 – in extracts. NH 4 + and NO 3 – in extracts were separated via a micro-diffusion method with magnesium oxide, Devarda’s alloy and oxalic acid absorption, and then the Isotope-Ratio Mass Spectrometry (IRMS) system (Europa Scientific Integra, Crewe, UK) was used to determine the 15 N abundances of NH 4 + and NO 3 – . The 15 N abundance of plant N was determined using the IRMS system with a Finnigan Flash EA 1112 elemental analyzer (Thermo Scientific, Bremen, Germany). The LI-6800 Portable Photosynthesis with a Multiphase Flash™ Fluorometer System (LI-COR Inc., Lincoln, NE, USA) was used to measure vegetable net photosynthetic rate, and the CO 2 concentration and light intensity of leaf chamber were set to 400 μmol mol −1 and 1800 μmol m −2 s −1 , respectively. Soil total microbial genomic DNA was extracted from 0.5 g fresh soil according to the manufacturer’s instructions of the Power Soil DNA Isolation Kit (Bioer Technology Co., Ltd). The real-time polymerase chain reaction quantification of genes associated to soil N cycling (including bacterial 16S, fungal ITS, archaeal and bacterial ammonia monooxygenase (AOA- amoA and AOB- amoA ), nitrite reductase ( nirK and nirS ), and nitrous oxide reductase ( nosZ )), and pathogens ( F.oxysporum , F.solani and Ralstonia solanacearum ) were performed on a CFX96TM Real-Time System (Bio-Rad Laboratories Inc., Hercules, CA, USA). Primer sets are listed in Table S1. The reaction mixtures were set according to Zhao et al. (2017), and standard curves were established according to Huang et al. (2015) and Zhao et al. (2017). The amplification efficiencies were 94–115%, with R 2 >0.99. The melting-curve analysis was used to evaluate amplification specificity, and copy numbers were subsequently calculated and converted to oven-dry basis. Two vegetable N uptake rates and ten soil gross N transformation rates were simultaneously quantified using the 15 N tracing tool Ntrace plant (Fig. S1) (He et al. 2020). The Markov Chain Monte Carlo algorithm is the key numerical technique for data analysis, and can determine the probability density function for each parameter and calculate true parameter uncertainties. The variance of the individual observations was taken into account in the misfit function between the modeled and observed data f (m) (Müller et al. 2007). The 15 N tracing model needed to be supplied with the concentrations and 15 N abundances of soil NH 4 + and NO 3 – , and plant N (mean ± standard deviation). The average values of twelve specific gross N transformation rates were calculated by simultaneously optimizing the kinetic parameters with setting as zero-order, first-order, or Michaelis-Menten kinetics, which could minimize the misfit between the observed and modeled values. The “mg N kg -1 soil d -1 ” was the unit for each gross N transformation rate. Based on Akaikes Information Criterion (AIC) (Cox et al. 2006) with MATLAB computing environment (Version 7, The MathWorks Inc.), the most suitable model was selected. For more details see He et al. (2020). Calculations and statistical analyses Soil net N mineralization ( net M ) and nitrification ( net N ) rates and NH 4 + retention time were calculated using the following equations: Net M = ( M + O Nrec ) – I TN Net N = N – I NO3 NH 4 + retention time = (NH 4 + addition + initial NH 4 + concentration) / ( I NH4 + O NH4 ) where the M was soil gross N transformation rates (i.e. mineralization of labile ( M Nlab ) and recalcitrant ( M Nrec ) organic N to NH 4 + ), O Nrec was heterotrophic nitrification (i.e. oxidation of recalcitrant organic N to NO 3 – ), O NH4 was autotrophic nitrification (i.e. oxidation of NH 4 + to NO 3 – ), N was total gross mineralization ( O Nrec + O NH4 ), I NH4 was microbial NH 4 + immobilization (i.e. immobilization of NH 4 + to N lab ( I NH4Nlab ) and N rec pool ( I NH4Nrec )), I NO3 was microbial NO 3 – immobilization (i.e. immobilization of NO 3 – to organic N pool), I TN was microbial N immobilization ( I NH4 + I NO3 ). T-test analysis was carried out to estimate the differences in soil N transformation rates, vegetable N uptake rates and the properties of soils and vegetables between RSD and CK, and presence and absence of vegetable, based on the averages, standard deviations and actual experimental repetitions (3 repetitions), using Sigma Plot (Version 3.5). Correlations between N transformation rates and soil properties were examined by spearman correlation analyses with two-sided tests (SPSS 23.0, Inc., USA). Results Soil and vegetable properties RSD significantly altered soil properties (Table 1). The soil pH was significantly higher in RSD treated soil than CK (7.27 vs. 5.81 for DST-RSD and DST-CK, 8.5 vs. 7.49 for DSC-RSD and DSC-CK). The EC value was reduced by RSD, from 430 to 371 µS cm − 1 in DST and 1256 to 387 µS cm − 1 in DSC. The concentrations of SOC, TN and DOC were 1.24−1.42, 1.05−1.09 and 1.09−4.70 times higher in RSD treatments compared to CK, respectively. The C/N ratio also significantly increased after RSD. There were no significant differences in soil TP and TK concentrations between RSD treatments and CK. After RSD, soil NO 3 − concentration decreased from 83 to 7 mg N kg − 1 soil in DST and 973 to 3 mg N kg − 1 soil in DSC, while soil NH 4 + concentration increased by 69- and 27- fold in DST and DSC, respectively. Soil AP and AK concentrations were significantly higher in RSD compared to CK, except for AP concentration in DST where no significant difference between RSD and CK was observed. The bacteria, AOA, AOB, nirK , nirS and nosZ abundances were significantly higher in RSD than CK, except in DST where there was no difference in nirK abundance between RSD and CK (Table 2 ). The fungi abundance in RSD was significantly lower compared to CK in DST, while in DSC it was not different from CK. The F. oxysporum , F. solani and Ralstonia solanacearum abundances were reduced significantly by RSD, except in DSC with no difference in Ralstonia solanacearum abundance between RSD and CK. After planting, soil pH generally decreased (i.e CC > CP and RC > RP; Fig. 1 a). The soil pH was significantly higher in RSD than CK, irrespectively of the presence of plants, i.e RC > CC and RP > CP. Plants significantly reduced soil DOC concentration in DST, while enhanced it in DSC (Fig. 1 b). Soil DOC concentration was significantly higher in CC than RC, while it was slightly higher in RP than CP but it was not significantly different in the two studied soils. After planting, no significant changes in the bacteria and AOA abundances were recorded (Fig. 1 c and e). RSD significantly enhanced the bacteria and AOA abundances (i.e. RC > CC and RP > CP). Bacterial abundance showed a positive correlation with soil pH (Fig. 3 a). In general, the presence of plants enhanced fungal abundance (Fig. 1 d). The fungal abundance in RSD was significantly lower compared to CK in DST (i.e. RC < CC and RP CC and RP > CP). The AOB abundance was significantly higher in CC than CP, and RC than RP (Fig. 1 f). RSD significantly enhanced the AOB abundance regardless of the presence of plants, i.e RC > CC and RP > CP. In DST, the vegetable biomass, net photosynthetic rate and N uptake were slightly higher in RP than CP, but not significantly (Fig. 1 g-i). In DSC, vegetable biomass, net photosynthetic rate and N uptake in RP were1.33 g pot − 1 , 17.09 µmol m − 1 s − 1 , and 46.93 mg N pot − 1 , respectively, and significantly higher than in CP (0.66 g pot − 1 , 6.29 µmol m − 1 s − 1 , and 26.78 mg N pot − 1 ). Gross and net N transformation rates The observed and modelled concentrations and 15 N abundances of NH 4 + , NO 3 − , and plant N matched well (R 2 > 0.94, Fig. S2-3). The 15 N- tracing analysis showed that soil gross N mineralization rates ( M ) were 1.4−3.9 times higher in RSD (RC and RP) than CK (CC and CP; Fig. 2 a). In DST, the presence of vegetables significantly reduced soil M (i.e. CP < CC and RP < CP). There was no difference in soil M between the presence and absence of vegetables in DSC, i.e CC ≈ CP and RC ≈ RP. Soil autotrophic nitrification rate ( O NH4 ) was significantly lower in CC than RC (3.89 vs. 7.95 mg N kg − 1 d − 1 in DST and 10.30 vs. 20.09 mg N kg − 1 d − 1 in DSC), and CP than RP (0.85 vs. 3.61 mg N kg − 1 d − 1 in DST and 1.85 vs. 3.15 mg N kg − 1 d − 1 in DSC; Fig. 2 b). The presence of vegetables significantly reduced soil O NH4 (i.e. CC > CP and RC > RP). Soil heterotrophic nitrification rate ( O Nrec ) were negligible in the absence of vegetables (i.e CC and RC), and increased to 0.81−2.61 mg N kg − 1 d − 1 in the presence of vegetables (i.e CP and RP). There were no differences in O Nrec between CC and RC, and between CP and RP in both soils, except in DSC where the O Nrec in RP was significantly higher than CP. O NH4 accounted for 30–100% of soil gross N nitrification rates ( N , O NH4 + O Nrec ). Microbial NH 4 + ( I NH4 ) and NO 3 − ( I NO3 ) immobilization rates ( I TN ) were negligible in the absence of vegetables (i.e CC and RC), while they significantly increased on average to 4.52 mg N kg − 1 d − 1 in the presence of vegetables (i.e CP and RP; Fig. 2 c). No significant differences in I NH4 and I NO3 were observed between RSD (RC) and CK (CC) in the absence of vegetables. In the presence of vegetables, the I NH4 and I NO3 in RSD (i.e. RP, on average 4.13 mg N kg − 1 d − 1 ) were significantly lower than CK (i.e. CP, on average 0.62 mg N kg − 1 d − 1 ). Plant NH 4 + uptake rate ( U NH4 ) in RSD (RP) was 2.29 and 13.49 mg N kg − 1 d − 1 in DST and DSC, respectively, and significantly higher than in CK (CP, 1.59 and 2.06 mg N kg − 1 d − 1 in DST and DSC, respectively; Fig. 2 d). Plant NO 3 − uptake rate ( U NO3 ) in RSD (RP) was not different from CK (CP) in DST, while it was significantly higher compared to CK in DSC (14.49 vs. 15.82 mg N kg − 1 d − 1 ). Soil net N mineralization ( Net M ) and nitrification ( Net N ) rates was 1.8–3.3 and 1.9–2.0 times higher, respectively, in RC than CC (Fig. 2 e and f). The ( U NH4 + I NH4 )/ M in RP was 9.2 and 1.1 folds lower in DST and DSC, respectively, compared to CP (Fig. 2 g). The presence of vegetables significantly enhanced the ( U NH4 + I NH4 )/ M (i.e. CC < CP and RC < RP). The ( O NH4 + I NH4 )/ M was significantly lower in RC than CC, and RP than CP in both soils, except in DSC where no significant difference in this ratio between CC and RC was observed (Fig. 2 h). The presence of plants significantly enhanced the ( O NH4 + I NH4 )/ M in DST (i.e. CC < CP, RC < RP), and generally reduced this ratio in DSC. The NH 4 + retention time ranged from 10.4 to 39.3 d in RC and RP, which was significantly higher than that in CC and CP, 4.6−13.1d; Fig. 2 i). NH 4 + retention time significantly higher in the presence of vegetables (i.e. CP and RP) than the absence of vegetables (i.e. CC and RC) in studied soils, except in DSC where the NH 4 + retention time was significantly lower in CP than CC. M was significantly and positively correlated with the bacterial abundance ( p < 0.05; Fig. 3 b) and soil pH ( p < 0.01; Fig. 3 c). O NH4 increased significantly with increasing soil pH ( p < 0.01; Fig. 3 d), and O NH4 showed a significantly negative correction with the ratio of ( U NH4 + I NH4 )/ M ( p < 0.01; Fig. 3 e). There was a significantly positive relationship between O Nrec and fungal abundance ( p < 0.05; Fig. 3 f). Discussion The results of the present study show that RSD significantly alters the gross N transformations and increases plant N uptake rates of soils that were used for continuous vegetable production. Consistent with our hypothesis, RSD significantly stimulated soil gross N mineralization and autotrophic nitrification rates, which enhanced the supply of N of both degraded soils and subsequent N uptake by plants. RSD improved soil quality (i.e. increasing soil pH, and decreasing EC and NO 3 − accumulation), further promoting vegetable N uptake, being in line with our hypotheses. RSD accelerated the production of soil NO 3 − Organic N mineralization to NH 4 + and subsequent oxidation to NO 3 − (autotrophic nitrification, O NH4 ) as well as direct oxidation of organic N to NO 3 − (heterotrophic nitrification, O Nrec ) were the two important pathways for NO 3 − production (Müller et al. 2011 ; Zhang et al. 2018 ). In this study, soil O NH4 was the major contributor producing NO 3 − , particularly in the absence of vegetables (Fig. 2 b). We found that RSD significantly enhanced O NH4 irrespectively of plants. Generally, soil N mineralization dominates NH 4 + production and is a key factor affecting O NH4 (Elrys et al. 2021b ; Schimel and Bennett 2004 ). In this study, Soil gross N mineralization ( M ) was significantly higher in RSD compared to CK and affected by plants (Fig. 2 a), indicating that RSD did stimulate soil M and subsequent NH 4 + production. This could be a main reason leading to higher soil NH 4 + concentration in RSD than CK (Table 1). It is widely reported that RSD can alter soil properties (Huang et al. 2016 ; Zhu et al. 2014 ), which is consistent with our results where soil pH, the concentrations of SOC, TN and DOC, and the bacteria abundance were enhanced by RSD (Table 1; Fig. 1 a, b and c). Numerous studies have shown that above-mentioned indicators were key for the regulation of M , with often significantly positive relationships (Elrys et al. 2021a ; Wang et al. 2016 ; Zhang et al. 2013 ). We confirmed this by a positive correlation of M with soil pH and the bacterial abundance (Fig. 4b and c). Furthermore, higher soil DOC, SOC and TN concentrations stimulated by RSD which in turn also affected soil pH and bacterial abundance led to increased M and therefore higher NH 4 + production. This is in line with the general observation that a higher NH 4 + production via M is also enhancing O NH4 , explaining the close correlation between O NH4 and M (Elrys et al. 2021b ). In addition, NH 4 + is also the N sources of vegetables and microorganisms in vegetable-soil systems (Hodge et al. 2000 ; Inselsbacher et al. 2010 ; Kuzyakov and Xu 2013 ). Therefore, vegetable NH 4 + uptake ( U NH4 ) and microbial NH 4 + immobilization ( I NH4 ) also affect O NH4 . Lower ( U NH4 + I NH4 )/ M ratio in RSD compared to CK can be caused by lower U NH4 and I NH4 or higher M , both enhancing the availability of NH 4 + to ammonia-oxidizing microorganisms which in turn stimulates O NH4 . This was supported by significantly negative relationships between O NH4 and ( U NH4 + I NH4 )/ M in this study (Fig. 3 e). Soil pH is a key factor regulating O NH4 (Elrys et al. 2021b ; Shammas 1986 ) and a significantly positive relationship between O NH4 and soil pH existed in this study (Fig. 3 c). Increasing soil pH can promote the release of NH 3 and enhance the activities of ammonia-oxidizing microorganisms (e.g. AOA and AOB), which has been shown to stimulate O NH4 (Li et al. 2018 ; Norton and Stark 2011 ; Stempfhuber et al. 2015 ). Here we provided evidence that soil pH and the AOA and AOB abundances were enhanced by RSD and in turn increased O NH4 . We also confirmed a stimulation of O Nrec by RSD as shown by Zhu et al. ( 2014 ), but only when plants were present. The exudates (e.g. acetic acids associated with O Nrec ) can be metabolized instantaneously and decomposed by 40% within two days (Laughlin et al. 2008 ) explaining for instance why O Nrec decreased from 0.34 to 0 mg N kg d − 1 at room temperature (He et al. 2020 ) and can explain negligible O Nrec in the absence of vegetables after 26 and 18 days cultivation. Surprisingly, RSD significantly enhanced O Nrec in the presence of vegetables (especially in DSC). After RSD treatment, soil NO 3 − content was reduced, and the NO 3 − production via O NH4 (ranging from 3.15 to 3.61 mg N kg − 1 d − 1 ) was not sufficient to meet plant NO 3 − uptake rates (ranging from 8.59 to 15.82 mg N kg − 1 d − 1 ). Thus, stimulation of O Nrec is one way to meet the N demand of the plants. These results indicate that O Nrec can be an important pathway for N supply to plants, which was in line with previous studies (Dan et al. 2022 ; He et al. 2022 ). Although both vegetables and microorganisms can assimilate soil NO 3 − , increasing soil NO 3 − production rates ( O NH4 + O Nrec ) due to RSD resulted in higher soil net nitrification rates compared to CK (positive relationship, Fig. 3 f), particularly in the absence of vegetables, which increased the leaching and accumulation risks of NO 3 − . This indicated that soil N fertilizer management after RSD treatment need to be carefully managed to avoid NO 3 − accumulation. RSD promoted vegetable N uptake by regulating soil gross N transformation and improving quality of degraded vegetable soils In this study, vegetable NH 4 + ( U NH4 ) and NO 3 − ( U NO3 ) uptake rates were generally higher in RP than CP (Fig. 2 d), indicating that RSD did promote vegetable N uptake. We found that significant differences in vegetable N uptake rates ( U TN , U NH4 + U NO3 ) between RP and CP were mainly attributed to differences in U NH4 . Although studied vegetables (i.e. tomato and cucumber) are characterized by NO 3 − -N preference, NH 4 + -N assimilation for vegetables is not only more energy efficient than NO 3 − -N, but also can promote root proliferation and keep the ionic balance (Al-Harbi 1995 ; Bloom et al. 2003 ; George et al. 2016 ; Nacry et al. 2013 ). Soil gross N mineralization ( M ), autotrophic nitrification ( O NH4 ) and NH 4 + immobilization ( O NH4 ) are key processes to govern soil NH 4 + concentration and NH 4 + retention time (Schimel and Bennett 2004 ), thereby affecting U NH4 . A lower ( O NH4 + I NH4 )/ M ratio in RP than CP causes a higher NH 4 + production (Fig. 2 h), leading to increased soil NH 4 + concentrations and in turn can promote U NH4 . In addition, compared to CP, longer NH 4 + retention time due higher soil initial NH 4 + concentration or lower O NH4 and I NH4 in RP (Fig. 2 g) can be an important factor responsible for higher U NH4 . Despite lower soil NO 3 − concentration in RSD compared to CK, U NO3 was generally higher in RP than CP (Fig. 2 d), which was likely duo to increasing soil gross nitrification rates after RSD. Higher soil gross nitrification rates in RP than CP (Fig. 2 b) can produce more NO 3 − for plant N uptake, explaining a stimulation of U NO3 . Clearly, RSD promoted vegetable N uptake by regulating soil gross N transformation processes in this study. In addition to soil N transformations, RSD can also alter soil physiochemical properties and thus influence the N uptake of plants. Soil physiochemical properties (e.g. soil pH, EC, microorganisms and structure) are important factors affecting vegetable growth (Bernstein 1975 ; Lau and Lennon 2012 ; Neina 2019 ; Passioura 1991 ). Long-term continuous cropping of the same or similar crop variety and high N fertilizer input can degrade the cultivated soil, e.g. soil NO 3 − accumulation, acidification and salinization, and increase of the abundances and diversities of soil pathogens, which can seriously affect plant growth (Guo et al. 2010 ; Hu et al. 2017 ; Huang et al. 2016 ; Shipton 1977 ). In this study, the quality of degraded soil was improved by RSD, i.e. increasing of soil pH and decreasing the contents of soil EC, NO 3 − and the abundances of pathogens ( F. oxysporum , F. solani and Ralstonia solanacearum ; Table 1), which supports plant growth, a higher net photosynthetic rate, total N uptake (Fig. 1 g-i and 2 d). Conclusions We were able to show that RSD significantly promotes plant N uptake, which can be attributed to two properties: 1) RSD decreased ( O NH4 + I NH4 )/ M , and increased NH 4 + retention time and production rates of soil NO 3 − , enhancing the N supply capacity of degraded soils; 2) RSD improved quality of degraded soil (i.e. increasing soil pH and decreasing the contents of soil EC and NO 3 − and pathogen abundances). Increasing soil net nitrification rates indicated that RSD accelerated soil NO 3 − production in the soil and increased the risk of leaching or accumulation of NO 3 − . Although RSD can improve the quality of degraded soils and promote plant N uptake, N fertilizer management after RSD still needs attention, otherwise NO 3 − accumulation or leaching will occur more rapidly. Declarations Acknowledgements This work was supported by the National Natural Science Foundation of China (grant number 41830642), the Postgraduate Research and Practice Innovation Program of Jiangsu Province (KYCX21_1332), and the “Double World-Classes”: Development in Geography Project. The study was carried out as part of the IAEA funded coordinated research project “Minimizing farming impacts on climate change by enhancing carbon and nitrogen capture and storage in Agro-Ecosystems (D1.50.16)” and was carried out in close collaboration with the German Science Foundation research unit DASIM (FOR 2337). Competing interests: None References Akhtar M (2000) Roles of organic soil amendments and soil organisms in the biological control of plant-parasitic nematodes: a review. 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Soil chemical properties in different treatments. Soils treatments pH EC SOC TN C/N TP TK DOC NH 4 + NO 3 - AP AK µS cm -1 g kg -1 g kg -1 mg kg -1 mg N kg -1 mg kg -1 DST CK 5.81±0.03b 430±6a 12.19±0.12b 1.61±0.06b 7.56±0.29b 1.40±0.06a 19±0a 151±3b 1±0b 83±1a 191±3a 260±4b RSD 7.27±0.00a 371±3b 15.16±0.12a 1.76±0.03a 8.60±0.15a 1.41±0.12a 19±1a 164±6a 69±1a 7±0b 190±5a 547±4a DSC CK 7.49±0.01b 1256±5a 15.12±0.47b 2.59±0.02b 5.83±0.18b 2.38±0.02a 22±0a 57±3b 6±08b 973±11a 97±4b 131±0b RSD 8.50±0.00a 387±16b 21.52±0.36a 2.71±0.03a 7.94±0.13a 2.38±0.05a 23±1a 268±16a 159±5a 3±0b 203±9a 507±6a DST, degraded tomato soil under tomato continuous cropping; DSC, degraded cucumber soil under cucumber continuous cropping; CK, a control (without reductive soil disinfestation); RSD, reductive soil disinfestation. EC, electrical conductivity; SOC, soil organic carbon; TN, total nitrogen; C/N, SOC: TN ratio; TP, total phosphorus; TK, total potassium; NH 4 + , ammonium; NO 3 – , nitrate; AP, available phosphorus; AK, available potassium. The different lowercase letters indicate the differences between CK and RSD in the same degraded soil ( p <0.05). Table 2. Soil microbial properties in different treatments. Soils treatments Bacteria Fungi AOA AOB nirK nirS nosZ F.o F.s R.s log 10 gene copies g -1 dry soil DST CK 9.88±0.04b 9.03±0.01a 7.00±0.05b 6.96±0.04b 7.86±0.16a 7.30±0.04b 7.92±0.04b 5.14±0.14a 6.17±0.09a 6.03±0.15a RSD 10.39±0.03a 8.75±0.05b 7.85±0.03a 7.97±0.03a 7.89±0.11a 8.33±0.02a 9.20±0.04a 4.48±0.13b 4.61±0.05b 5.48±0.13b DSC CK 10.17±0.06b 8.29±0.08a 8.15±0.03b 6.52±0.10b 7.01±0.12b 7.38±0.08b 7.52±0.07b 5.14±0.19a 4.67±0.10a 5.72±0.19a RSD 10.85±0.05a 8.31±0.04a 8.35±0.01a 6.98±0.05a 7.84±0.15a 8.24±0.09a 8.64±0.10a 4.81±0.05b 4.11±0.09b 5.79±0.07a DST, degraded tomato soil under tomato continuous cropping; DSC, degraded cucumber soil under cucumber continuous cropping; CK, a control (without reductive soil disinfestation); RSD, reductive soil disinfestation. AOA, ammonia-oxidizing archaea; AOB, ammonia-oxidizing bacteria; nirK and nirS , the genes of nitrite reductase; nosZ , the gene of nitrous oxide reductase. The different lowercase letters indicate the differences between CK and RSD in the same degraded vegetable soil ( p <0.05). Supplementary Files SupportingMaterialsCM.docx Cite Share Download PDF Status: Under Review Version 1 posted Editorial decision: Minor revisions 24 Jul, 2022 Reviewers agreed at journal 04 Jul, 2022 Reviewers invited by journal 04 Jul, 2022 Editor invited by journal 01 Jul, 2022 Editor assigned by journal 01 Jul, 2022 First submitted to journal 28 Jun, 2022 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. 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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-1805779","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":118522530,"identity":"ace8f1f4-dbfa-4c45-b422-faedb55aa709","order_by":0,"name":"jinbo zhang","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA8klEQVRIiWNgGAWjYDACCRDBI8FgwMB8AMJhSCBaC1sCKVqAwICBxwDKJKBFfnbzs4dfZCzyzNl7vj2w3HGYgZ89x4Dh5w7cWhjnHDM3luGRKLbsObvdQPLMYQbJnjcGjL1ncGthlkgwk5bgkUjccCN3m4Rk22EGgxs5BsyMbbi1sEmkf4NqyXkG1mJPSAuPRI6Z5AeIFjaILRIEtEhI5JRJAzUm7uw5ZgbUks4jceZZwcFePFrkZ6Rvk/zZU5e4nb35mbRkm7Ucf3vyxgc/8WgBBwFvDywsgC4FMQ7g1wAM6B8/oIwPhJSOglEwCkbBiAQAPfRJ3TzkWe0AAAAASUVORK5CYII=","orcid":"https://orcid.org/0000-0002-5659-7921","institution":"School of Geography Sciences, Nanjing Normal University","correspondingAuthor":true,"submittingAuthor":false,"prefix":"","firstName":"jinbo","middleName":"","lastName":"zhang","suffix":""},{"id":118522531,"identity":"32b9339d-6a4b-4a6f-b4be-8a3a0cd92c25","order_by":1,"name":"Xiaoqian Dan","email":"","orcid":"","institution":"","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Xiaoqian","middleName":"","lastName":"Dan","suffix":""},{"id":118522532,"identity":"671bfcbd-8a89-48d9-ac3e-43977306bc20","order_by":2,"name":"Mengqiu He","email":"","orcid":"","institution":"","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Mengqiu","middleName":"","lastName":"He","suffix":""},{"id":118522533,"identity":"ef9faaa0-16f5-46c3-9e58-8f957ea7fea1","order_by":3,"name":"Shending Chen","email":"","orcid":"","institution":"","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Shending","middleName":"","lastName":"Chen","suffix":""},{"id":118522534,"identity":"943a5ebd-1eb7-49e4-9004-241af091b400","order_by":4,"name":"Xiaoxiang He","email":"","orcid":"","institution":"","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Xiaoxiang","middleName":"","lastName":"He","suffix":""},{"id":118522535,"identity":"bf2e48fa-07f1-48e3-b704-b8bbdbad4ed4","order_by":5,"name":"Chang Zhao","email":"","orcid":"","institution":"","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Chang","middleName":"","lastName":"Zhao","suffix":""},{"id":118522536,"identity":"cc523b9f-7a99-4bae-87a1-9e7a1bb54b9e","order_by":6,"name":"Lei Meng","email":"","orcid":"","institution":"","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Lei","middleName":"","lastName":"Meng","suffix":""},{"id":118522537,"identity":"afca23ed-c8f7-435f-b545-1bcb36d2f3b9","order_by":7,"name":"Zucong Cai","email":"","orcid":"","institution":"","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Zucong","middleName":"","lastName":"Cai","suffix":""},{"id":118522538,"identity":"70ff2aab-32c0-4dd3-b663-1a2948a9908e","order_by":8,"name":"Christoph Müller","email":"","orcid":"","institution":"","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Christoph","middleName":"","lastName":"Müller","suffix":""}],"badges":[],"createdAt":"2022-06-29 02:37:19","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-1805779/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-1805779/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":23773207,"identity":"382fbb17-5556-4557-a83f-f92957038945","added_by":"auto","created_at":"2022-07-12 16:36:21","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":135558,"visible":true,"origin":"","legend":"\u003cp\u003eSoil and vegetable properties in different treatments after planting.\u003c/p\u003e\u003cp\u003eDST, degraded soil under tomato continuous cropping; DSC, degraded soil under continuous cucumber cropping. CC, control (without reductive soil disinfestation) without plants; RC, reductive soil disinfestation without plantd; CP, control (without reductive soil disinfestation) with plants; RP, reductive soil disinfestation with plants. DOC, soil dissolved organic carbon; AOA, ammonia-oxidizing archaea; AOB, ammonia-oxidizing bacteria. Vertical bars indicate standard deviations of the mean. Different lowercase letters represent significant differences between CK and RSD, and different capital letters represent significant differences between the absence and presence of plants (\u003cem\u003ep\u003c/em\u003e \u0026lt;0.05).\u003c/p\u003e","description":"","filename":"Fig1.jpg","url":"https://assets-eu.researchsquare.com/files/rs-1805779/v1/2b2c0cbe915dea3a01cca44b.jpg"},{"id":23773208,"identity":"d7ddfb16-db3c-47ee-aa48-5d0ea292e4b1","added_by":"auto","created_at":"2022-07-12 16:36:21","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":114730,"visible":true,"origin":"","legend":"\u003cp\u003eGross and net N transformation rates in different treatments.\u003c/p\u003e\u003cp\u003eDST, degraded soil under tomato continuous cropping; DSC, degraded soil under continuous cucumber cultivation. CC, control (without reductive soil disinfestation) without plants; RC, reductive soil disinfestation without plants; CP, control (without reductive soil disinfestation) with plants; RP, reductive soil disinfestation with plants. \u003cem\u003eM\u003c/em\u003e, total gross N mineralization rates;\u003cem\u003e ONH4\u003c/em\u003e, oxidation rate of NH4+ to NO3– (autotrophic nitrification rate); \u003cem\u003eONrec\u003c/em\u003e, oxidation rate of recalcitrant organic N pool to NO3– (heterotrophic nitrification rate); \u003cem\u003eN\u003c/em\u003e, total gross N nitrification rates;\u003cem\u003e INH4\u003c/em\u003e, total gross NH4+ immobilization rate; \u003cem\u003eINO3\u003c/em\u003e, immobilization rate of NO3– to organic N pool; \u003cem\u003eITN\u003c/em\u003e, total gross N immobilization rates (\u003cem\u003eINH4\u003c/em\u003e + \u003cem\u003eINO3\u003c/em\u003e); \u003cem\u003eUNH4\u003c/em\u003e, vegetable NH4+ uptake rate; \u003cem\u003eUNO3\u003c/em\u003e, vegetable NO3– uptake rate; \u003cem\u003eUTN\u003c/em\u003e, vegetable total N uptake rates (\u003cem\u003eUNH4\u003c/em\u003e + \u003cem\u003eUNO3\u003c/em\u003e); \u003cem\u003eNet M\u003c/em\u003e, net N mineralization rate; \u003cem\u003eNet N\u003c/em\u003e, net nitrification rate. Vertical bars represent standard deviations of the mean. Different lowercase letters represent significant differences between CK and RSD, and different capital letters represent significant differences between the absence and presence of vegetable (\u003cem\u003ep\u003c/em\u003e \u0026lt;0.05).\u003c/p\u003e","description":"","filename":"Fig2.jpg","url":"https://assets-eu.researchsquare.com/files/rs-1805779/v1/023493a589dcd52c8076dd2b.jpg"},{"id":23774121,"identity":"029146ac-0fa0-4dac-a370-fd6ac6948f6e","added_by":"auto","created_at":"2022-07-12 16:41:21","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":113524,"visible":true,"origin":"","legend":"\u003cp\u003eThe relationships among soil gross N transformation rates and soil properties.\u003c/p\u003e\u003cp\u003e\u003cem\u003eM\u003c/em\u003e, total gross N mineralization rates;\u003cem\u003e ONH4\u003c/em\u003e, oxidation rate of NH4+ to NO3– (autotrophic nitrification rate); \u003cem\u003eONrec\u003c/em\u003e, oxidation rate of recalcitrant organic N pool to NO3– (heterotrophic nitrification rate); \u003cem\u003eN\u003c/em\u003e, total gross N nitrification rates;\u003cem\u003e INH4\u003c/em\u003e, total gross NH4+ immobilization rate; \u003cem\u003eUNH4\u003c/em\u003e, vegetable NH4+ uptake rate. Points represent average rates, and dashed lines represent the 95% confidence interval of the regression line. Vertical bars represent standard deviations of the mean.\u003c/p\u003e","description":"","filename":"Fig3.jpg","url":"https://assets-eu.researchsquare.com/files/rs-1805779/v1/c88d59164cd4eeb74e0888bf.jpg"},{"id":23774122,"identity":"1fa7eb38-2e1d-4cd4-b2fc-757978b57764","added_by":"auto","created_at":"2022-07-12 16:41:24","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":582622,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-1805779/v1/d8c65497-18f0-4e41-9cf0-0b71c89a5636.pdf"},{"id":23773210,"identity":"67891289-05d3-4a87-a838-b224ba626827","added_by":"auto","created_at":"2022-07-12 16:36:21","extension":"docx","order_by":6,"title":"","display":"","copyAsset":false,"role":"supplement","size":287061,"visible":true,"origin":"","legend":"","description":"","filename":"SupportingMaterialsCM.docx","url":"https://assets-eu.researchsquare.com/files/rs-1805779/v1/c1eab4b8cd68d030dca663de.docx"}],"financialInterests":"","formattedTitle":"Reductive soil disinfestation promoted vegetable N uptake by regulating soil gross N transformations and improving the quality of degraded vegetable soils","fulltext":[{"header":"Introduction","content":"\u003cp\u003eDue to the high demand in the markets, the greenhouse vegetable production (GVP) is increasing rapidly worldwide. By 2018, the global area of GVP was approximately 5.6\u0026nbsp;million ha (Cuesta \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Fei et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). For high yield and economic profit, agricultural practices, such as intensive crop rotations (Jia et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2012\u003c/span\u003e) and high rates of N fertilizer application (often exceeding 1000 kg N ha-1, Ju et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Shi et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2009\u003c/span\u003e), are widespread, especially in GVP. However, low N absorption capacity arising from the sparse and shallow root systems and short growth cycle of vegetables induce low N use efficiency (NUE) in GVP, which is generally\u0026thinsp;\u0026lt;\u0026thinsp;20% (Ren et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Ti et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Consequently, long-term GVP lead to soil degradation, e.g. accumulation of NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e in the soil, acidification, salinization and severe soil-borne diseases (Guo et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Hu et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Huang et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). This deterioration in the quality, safety and yield of vegetables led to environment problems and jeopardized the sustainable development of GVP (Albornoz \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Hu et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eReductive soil disinfestation (RSD), as an environmentally friendly method developed in Japan and Netherlands in 2000 (Blok et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Shinmura \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2000\u003c/span\u003e), has been widely promoted to improve the quality of degraded vegetable soils (Butler et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2012b\u003c/span\u003e; Huang et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Li et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Briefly, in the RSD method, easily decomposable organic material (e.g. molasses, alfalfa, maize straw) is homogeneously added into the soil. The soil is then irrigated until saturation, covered with plastic film and cultivated at a temperature of 30 to 35\u0026deg;C for 2\u0026minus;4 weeks. This treatment creates strong reductive conditions and high temperatures to effectively improve the physical, chemical and biological properties of degraded vegetable soils (Butler et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2012b\u003c/span\u003e; Huang et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Liu et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). For instance, the addition of organic matter can improve soil structure (such as enhancements of soil holding water and ions exchange), in favor of vegetable growth (Akhtar and Malik 2000; Oka \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Combining strong reductive condition with easily decomposable organic matter amendment can simulate the growth of soil denitrification microorganisms (e.g. \u003cem\u003enirK\u003c/em\u003e, \u003cem\u003enirS\u003c/em\u003e and \u003cem\u003enosZ\u003c/em\u003e) (Huang et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Huang et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Zhao et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), which can promote the denitrification of NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e and further rapidly remove soil accumulated NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e. In addition, RSD can decrease soil electrical conductivity (EC), enhance soil pH and NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e content, and efficiently suppress soil-borne plant pathogens and plant-parasitic nematodes (Butler et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2012a\u003c/span\u003e; Huang et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Liu et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Zhao et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Thus, degraded vegetable soils treated with RSD can provide a more favorable growing environment for vegetables and improve their N uptake, which promotes NUE and vegetable yield and reduces the environmental risk of N loss.\u003c/p\u003e \u003cp\u003eIn addition, the interactions between soil gross N transformations could affect plant N uptake, as they determine the amount and form of available N in the soil (Zhang et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). It is widely reported that RSD significantly alters soil properties (e.g. soil pH, total N, organic carbon (C), NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e, NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e, AOB, AOA, bacteria and fungi) (Huang et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Zhu et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2014\u003c/span\u003e), which were key factors regulating soil gross N transformations. For instance, soil organic substrates (such total N and C) were vital energy sources for microorganisms and could drive microbial-mediated soil N cycling (e.g. soil gross N mineralization and heterotrophic nitrification) (Elrys et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2021a\u003c/span\u003e; Elrys et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2021b\u003c/span\u003e; Zhang et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Zhang et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Increasing soil pH can stimulate autotrophic nitrification by enhancing the release of NH\u003csub\u003e3\u003c/sub\u003e and the activities of ammonia-oxidizing microorganisms (e.g. AOA and AOB) (Li et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Norton and Stark \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Stempfhuber et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). These were confirmed by Zhu et al. (\u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2014\u003c/span\u003e) where RSD significantly enhanced heterotrophic nitrification and autotrophic nitrification by increasing soil pH, total N and organic carbon. In addition, the increase of soil microbial activity resulted in higher soil microbial immobilization rates after RSD. Although Zhu et al. (\u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2014\u003c/span\u003e) concluded that RSD altered degraded vegetable soils gross N transformation rates. However, their results lacked the influence of vegetables N uptake, leading to an incomplete understanding of soil N transformations after RSD. Soil-plant feedbacks influence soil gross N transformations through transfer of organic matter into the soil in the form of litter or root exudates, which reduces the substrates of N transformations and affects the activity of microorganisms involved in N transformations (Dan et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; He et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Nacry et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Zhao et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). At present, the effects of RSD on the interactions between vegetable N uptake and soil gross N transformation rates are still unclear.\u003c/p\u003e \u003cp\u003eA series of \u003csup\u003e15\u003c/sup\u003eN tracing pot experiments were carried out with vegetables to quantify the effects of RSD treatment on two degraded soils in terms of soil gross N transformation rates and plant N uptake rates using a \u003csup\u003e15\u003c/sup\u003eN tracing approach (\u003cem\u003eNtrace\u003c/em\u003e\u003csub\u003eplant\u003c/sub\u003e). Our study was carried out to test the following hypotheses: 1) soil gross N mineralization and autotrophic nitrification can be stimulated by RSD, considering that soil total N, pH and activities of ammonia-oxidizing microorganisms increased after RSD; 2) RSD can promote vegetable N uptake through effectively improving the quality of degraded vegetable soil.\u003c/p\u003e"},{"header":"Material And Methods","content":"\u003cp\u003e\u003cem\u003eSoil samples\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eDegraded greenhouse soils that had been under continuous cropping with tomato (Lycopersicon\u003cem\u003e\u0026nbsp;esculentum\u0026nbsp;\u003c/em\u003eL.) (DST) and cucumber (\u003cem\u003eCucumis sativus\u0026nbsp;\u003c/em\u003eL.) (DSC)\u0026nbsp;were sampled from greenhouses in Nanjing, Jiangsu Province, China. Soils were classified as udalfs (Alfisols) based\u0026nbsp;\u003ca href=\"javascript%3A;\"\u003eto\u003c/a\u003e the soil taxonomy of the United States of America. On DST, tomato and strawberry had been grown continuously for more than ten years, leading to problems with continuous cultivation of tomatoes and strawberries. On DSC, more than 20 crops of continuous monoculture cultivation which resulted in an obstacle to continuous cucumber cultivation. Both two soils were collected after vegetable harvest. At each greenhouse, 4 plots with 1 m \u0026times;1 m were randomly selected, and surface soil (0-20 cm) was sampled and then mixed in equal amounts, subsequently sieved (2 mm) and roots were removed. Each soil was divided into two parts, one was treated by the reductive soil disinfestation (RSD) method, the other was store at 4\u0026nbsp;\u0026deg;C and served as a control (CK). Therefore, four soil samples, i.e. DST without and with RSD (named as DST-CK and DST-RSD, respectively) and DSC without and with RSD (named as DSC-CK and DSC-RSD, respectively), were used for the experiments. For RSD method, each soil was incorporated with alfalfa meal at a ratio of 1/50 (dry soil/alfalfa meal, \u003cem\u003ew\u003c/em\u003e/\u003cem\u003ew\u003c/em\u003e), irrigated to 100% water hold capacity (WHC) and then evenly mixed. Treated soils were packed in zip-lock bags, sealed, placed in an incubator with 30\u0026nbsp;\u0026deg;C and cultivated for 21 days. Then, soils were dried to approximate 20% WHC, and passed through a 2 mm sieve. Each soil sample (whether RSD or not) was divided into 3 subsamples to determine the soil chemical (Table1), microbial properties (Table2) and conduct \u003csup\u003e15\u003c/sup\u003eN-tracing studies, respectively. \u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e\u003csup\u003e15\u003c/sup\u003e\u003c/em\u003e\u003cem\u003eN-tracing experiments\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eEight treatments were setup, based on the four soil samples (e.g. DST-CK, DST-RSD, DSC-CK and DSC-RSD ) with and without plants, i.e. 1) only DST-CK soil (DST-CC), 2) DST-CK soil with tomato (DST-CP), 3) only DST-RSD soil (DST-RC), 4) DST-RSD soil with tomato (DST-RP), 5) only DSC-CK soil (DSC-CC), 6) DSC-CK soil with cucumber (DSC-CP), 7) only\u0026nbsp;DSC-RSD\u0026nbsp;soil (DSC-RC), and 8)\u0026nbsp;DSC-RSD\u0026nbsp;soil with cucumber (DSC-RP).\u003c/p\u003e\n\u003cp\u003eTomato and cucumber seeds were\u0026nbsp;sterilized\u0026nbsp;using 5% alcohol for 5 min, and washed 4 times using deionized water. The seeds were germinated for 2-3 days on petri dishes (with a moistened gauze on the bottom) placing in a 30\u003csup\u003e\u0026nbsp;\u0026deg;\u003c/sup\u003eC-incubator. After germination the seedlings with uniform growth rate were selected and transplanted to the seeding plug tray. When its first true leaf had been fully developed, seedlings of uniform growth were transplanted into 360-mL pots containing 0.2 kg soil (oven-dry basis) after the nutritional material on roots was largely removed.\u0026nbsp;Meanwhile, the treatments without planting were also set up. Each pot received nutrient solutions containing N, K and P with 50, 30 and 20 mg kg\u003csup\u003e-1\u003c/sup\u003e soil, respectively, and trace elements, and was placed in a greenhouse for cultivation. The soil water content maintained at approximately 60% WHC by daily water additions.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eAfter\u0026nbsp;26 and 18 days\u0026nbsp;of transplanting tomato and cucumber seedlings, respectively, a \u003csup\u003e15\u003c/sup\u003eN tracing experiment was carried out including all treatments. Two \u003csup\u003e15\u003c/sup\u003eN-labeled groups were established for each treatment, where either NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e was \u003csup\u003e15\u003c/sup\u003eN labelled (\u003csup\u003e15\u003c/sup\u003eNH\u003csub\u003e4\u003c/sub\u003eNO\u003csub\u003e3\u003c/sub\u003e, 10.20 atom% for all treatments) or where\u0026nbsp;NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026ndash;\u003c/sup\u003e was \u003csup\u003e15\u003c/sup\u003eN labelled\u0026nbsp;(NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e15\u003c/sup\u003eNO\u003csub\u003e3\u003c/sub\u003e, 10.12 atom% for\u0026nbsp;DST-CC, DST-CP, DST-RC, DST-RP,\u0026nbsp;DSC-RC,\u0026nbsp;DSC-RP treatments, and 99.23 atom% for DSC-CC, DSC-CP\u0026nbsp;treatments). 2-mL each \u003csup\u003e15\u003c/sup\u003eN solution was uniformly applied\u0026nbsp;to each pot using a 4-needle injection technique\u0026nbsp;at a\u0026nbsp;rate of 30 mg NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e-N kg\u003csup\u003e-1\u003c/sup\u003e soil and 30 mg\u0026nbsp;NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026ndash;\u003c/sup\u003e-N kg\u003csup\u003e-1\u003c/sup\u003e soil for DST, and 50 mg NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e-N kg\u003csup\u003e-1\u003c/sup\u003e soil and 50 mg\u0026nbsp;NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026ndash;\u003c/sup\u003e-N kg\u003csup\u003e-1\u003c/sup\u003e soil (He et al. 2022). At 0.5, 24, 48 and 72 h after \u003csup\u003e15\u003c/sup\u003eN labelling, both soil and plants were collected. 1 M potassium chloride\u0026nbsp;(KCl) solution (soil/solution with 1/5) was used to extract NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e and\u0026nbsp;NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026ndash;\u003c/sup\u003e in soil for determination of concentrations and \u003csup\u003e15\u003c/sup\u003eN abundances of NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e and\u0026nbsp;NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026ndash;\u003c/sup\u003e. Other soil properties, i.e. Soil pH, DOC (dissolved organic carbon), the bacteria, fungi, AOA (ammonia-oxidizing archaea) and AOB (ammonia-oxidizing bacteria) abundances, were also measured. Residual inorganic N on plant surfaces were thoroughly rinsed with 1 M KCl solution and then deionized water. After that, vegetables (including root) were\u0026nbsp;de-enzymed in ovens with 105\u0026nbsp;\u0026deg;C for 0.5 h, dried in ovens with 80 \u003csup\u003eo\u003c/sup\u003eC to a constant weight.\u0026nbsp;Dried vegetables were weighed to obtain their biomass, and then grinded\u0026nbsp;and sieved (0.15 mm) to analyze their N concentrations and \u003csup\u003e15\u003c/sup\u003eN abundances.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eAnalytical methods\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eSoil pH (soil/water, 2.5:1) and EC (electrical conductivity) (soil/water, 5:1) were measured using DMP-2mV/pH detector (Quark Ltd., Nanjing, China) and conductivity detector (Kang Yi Crops., Nanjing, China), respectively. Soil organic carbon (SOC) and total N (TN) were analyzed by wet digestion with H\u003csub\u003e2\u003c/sub\u003eSO\u003csub\u003e4\u003c/sub\u003e-K\u003csub\u003e2\u003c/sub\u003eCr\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e7\u003c/sub\u003e and semi-micro Kjeldahl digestion with Se, CuSO\u003csub\u003e4\u003c/sub\u003e, and K\u003csub\u003e2\u003c/sub\u003eSO\u003csub\u003e4\u003c/sub\u003e as catalysts, respectively. Soil dissolved organic C (DOC) content was determined using the Analyzer Multi N/C (Analytic Jena, Jena, Germany). NaOH fusion and Mo-Sb colorimetry, NaOH fusion and flame photometer, NaHCO\u003csub\u003e3\u003c/sub\u003e digestion and Mo-Sb colorimetry, as well as\u0026nbsp;CH\u003csub\u003e3\u003c/sub\u003eCOONH\u003csub\u003e4\u003c/sub\u003e digestion and flame photometer were used to analyze the total soil phosphorus (TP), total potassium (TK), available phosphorus (AP) and available potassium (AK), respectively, according to Bao (2000). Soils were extracted with 1 M KCl solution (soil to solution ratio, 1:5), the automated continuous flow wet chemistry analyzer (SA1000, Skalar, Netherlands) was used to analyze the concentrations of NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e and\u0026nbsp;NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026ndash;\u003c/sup\u003e in extracts. NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e and\u0026nbsp;NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026ndash;\u003c/sup\u003e in extracts were separated via a micro-diffusion method with magnesium oxide, Devarda\u0026rsquo;s alloy and oxalic acid absorption, and then the Isotope-Ratio Mass Spectrometry (IRMS) system (Europa Scientific Integra, Crewe, UK) was used to determine the\u0026nbsp;\u003csup\u003e15\u003c/sup\u003eN abundances of\u0026nbsp;NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e and\u0026nbsp;NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026ndash;\u003c/sup\u003e.\u0026nbsp;The\u0026nbsp;\u003csup\u003e15\u003c/sup\u003eN abundance of\u0026nbsp;plant\u0026nbsp;N was determined using the IRMS system with a Finnigan Flash EA 1112 elemental analyzer (Thermo Scientific, Bremen, Germany).\u0026nbsp;The LI-6800 Portable Photosynthesis with a Multiphase Flash\u0026trade; Fluorometer System (LI-COR Inc., Lincoln, NE, USA) was used to measure vegetable net photosynthetic rate, and the CO\u003csub\u003e2\u003c/sub\u003e concentration and light intensity of leaf chamber were set to 400 \u0026mu;mol mol\u003csup\u003e\u0026minus;1\u003c/sup\u003e and 1800 \u0026mu;mol m\u003csup\u003e\u0026minus;2\u003c/sup\u003e s\u003csup\u003e\u0026minus;1\u003c/sup\u003e, respectively.\u003c/p\u003e\n\u003cp\u003eSoil total microbial genomic DNA was\u0026nbsp;extracted from 0.5 g fresh soil according to the manufacturer\u0026rsquo;s instructions of the Power Soil DNA Isolation Kit (Bioer Technology Co., Ltd). The real-time polymerase chain reaction quantification of genes associated to soil N cycling (including bacterial 16S, fungal ITS, archaeal and bacterial ammonia monooxygenase (AOA-\u003cem\u003eamoA\u003c/em\u003e and AOB-\u003cem\u003eamoA\u003c/em\u003e), nitrite reductase (\u003cem\u003enirK\u003c/em\u003e and \u003cem\u003enirS\u003c/em\u003e), and nitrous oxide reductase (\u003cem\u003enosZ\u003c/em\u003e)), and\u0026nbsp;pathogens (\u003cem\u003eF.oxysporum\u0026nbsp;\u003c/em\u003e, \u003cem\u003eF.solani\u003c/em\u003e and\u0026nbsp;\u003cem\u003eRalstonia solanacearum\u003c/em\u003e) were performed on a CFX96TM Real-Time System (Bio-Rad Laboratories Inc., Hercules, CA, USA). Primer sets are listed in Table S1. The reaction mixtures were set according to Zhao et al. (2017), and standard curves were established according to Huang et al. (2015) and Zhao et al. (2017). The amplification efficiencies were 94\u0026ndash;115%, with R\u003csup\u003e2\u0026nbsp;\u003c/sup\u003e\u0026gt;0.99. The melting-curve analysis was used to evaluate amplification specificity, and copy numbers were subsequently calculated and converted to oven-dry basis.\u003c/p\u003e\n\u003cp\u003eTwo vegetable\u0026nbsp;N uptake rates and ten soil gross N transformation rates were simultaneously quantified using the \u003csup\u003e15\u003c/sup\u003eN tracing tool \u003cem\u003eNtrace\u003c/em\u003e\u003csub\u003eplant\u003c/sub\u003e (Fig. S1) (He et al. 2020). The Markov Chain Monte Carlo algorithm is the key numerical technique for data analysis, and can determine the probability density function for each parameter and calculate true parameter uncertainties. The variance of the individual observations was taken into account in the misfit function between the modeled and observed data \u003cem\u003ef\u003c/em\u003e(m) (M\u0026uuml;ller et al. 2007). The \u003csup\u003e15\u003c/sup\u003eN tracing model needed to be supplied with the concentrations and \u003csup\u003e15\u003c/sup\u003eN abundances of soil NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e and NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026ndash;\u003c/sup\u003e, and plant N (mean \u0026plusmn; standard deviation). The average values of twelve specific gross N transformation rates were calculated by simultaneously optimizing the kinetic parameters with setting as zero-order, first-order, or Michaelis-Menten kinetics, which could minimize the misfit between the observed and modeled values. The \u0026ldquo;mg N kg\u003csup\u003e-1\u003c/sup\u003e soil d\u003csup\u003e-1\u003c/sup\u003e\u0026rdquo; was the unit for each gross N transformation rate. Based on Akaikes Information Criterion (AIC) (Cox et al. 2006) with MATLAB computing environment (Version 7, The MathWorks Inc.), the most suitable model was selected. For more details see He et al. (2020).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eCalculations and statistical analyses\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eSoil net N mineralization (\u003cem\u003enet M\u003c/em\u003e) and nitrification (\u003cem\u003enet N\u003c/em\u003e)\u0026nbsp;rates and NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e retention time were calculated using the following equations:\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eNet M\u003c/em\u003e = (\u003cem\u003eM\u003c/em\u003e + \u003cem\u003eO\u003csub\u003eNrec\u003c/sub\u003e\u003c/em\u003e) \u0026ndash;\u0026nbsp;\u003cem\u003eI\u003csub\u003eTN\u003c/sub\u003e\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eNet N\u003c/em\u003e = \u003cem\u003eN\u003c/em\u003e \u0026ndash;\u0026nbsp;\u003cem\u003eI\u003csub\u003eNO3\u003c/sub\u003e\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e retention time = (NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e addition + initial NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u0026nbsp;\u003c/sup\u003econcentration) / (\u003cem\u003eI\u003csub\u003eNH4\u003c/sub\u003e\u003c/em\u003e +\u0026nbsp;\u003cem\u003eO\u003csub\u003eNH4\u003c/sub\u003e\u003c/em\u003e)\u003c/p\u003e\n\u003cp\u003ewhere the \u003cem\u003eM\u003c/em\u003e was soil gross N transformation rates\u0026nbsp;(i.e. mineralization of labile (\u003cem\u003eM\u003csub\u003eNlab\u003c/sub\u003e\u003c/em\u003e) and recalcitrant (\u003cem\u003eM\u003csub\u003eNrec\u003c/sub\u003e\u003c/em\u003e) organic N to NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e), \u003cem\u003eO\u003csub\u003eNrec\u003c/sub\u003e\u003c/em\u003e was heterotrophic nitrification (i.e. oxidation of recalcitrant organic N to NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026ndash;\u003c/sup\u003e), \u003cem\u003eO\u003csub\u003eNH4\u003c/sub\u003e\u003c/em\u003e was\u0026nbsp;autotrophic nitrification (i.e. oxidation of NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e to NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026ndash;\u003c/sup\u003e), \u003cem\u003eN\u003c/em\u003e was total gross mineralization (\u003cem\u003eO\u003csub\u003eNrec\u003c/sub\u003e\u0026nbsp;\u003c/em\u003e+\u0026nbsp;\u003cem\u003eO\u003csub\u003eNH4\u003c/sub\u003e\u003c/em\u003e),\u003cem\u003e\u0026nbsp;I\u003csub\u003eNH4\u003c/sub\u003e\u003c/em\u003e was microbial\u0026nbsp;NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e immobilization (i.e. immobilization of NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e to \u003cem\u003eN\u003csub\u003elab\u003c/sub\u003e\u0026nbsp;\u003c/em\u003e(\u003cem\u003eI\u003csub\u003eNH4Nlab\u003c/sub\u003e\u003c/em\u003e) and \u003cem\u003eN\u003csub\u003erec\u003c/sub\u003e\u003c/em\u003e pool\u0026nbsp;(\u003cem\u003eI\u003csub\u003eNH4Nrec\u003c/sub\u003e\u003c/em\u003e)),\u003cem\u003e\u0026nbsp;I\u003csub\u003eNO3\u003c/sub\u003e\u003c/em\u003e was microbial\u0026nbsp;NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026ndash;\u003c/sup\u003e immobilization (i.e. immobilization of NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026ndash;\u003c/sup\u003e to organic N pool), \u003cem\u003eI\u003csub\u003eTN\u003c/sub\u003e\u003c/em\u003e was microbial\u0026nbsp;N immobilization (\u003cem\u003eI\u003csub\u003eNH4\u003c/sub\u003e\u003c/em\u003e + \u003cem\u003eI\u003csub\u003eNO3\u003c/sub\u003e\u003c/em\u003e).\u003c/p\u003e\n\u003cp\u003eT-test analysis was carried out to estimate the differences in soil N transformation rates, vegetable N uptake rates and the properties of soils and vegetables between RSD and CK, and presence and absence of vegetable, based on the averages, standard deviations and actual experimental repetitions (3 repetitions), using Sigma Plot (Version 3.5). Correlations between N transformation rates and soil properties were examined by spearman correlation analyses with two-sided tests (SPSS 23.0, Inc., USA).\u003c/p\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003eSoil and vegetable properties\u003c/h2\u003e \u003cp\u003eRSD significantly altered soil properties (Table\u0026nbsp;1). The soil pH was significantly higher in RSD treated soil than CK (7.27 vs. 5.81 for DST-RSD and DST-CK, 8.5 vs. 7.49 for DSC-RSD and DSC-CK). The EC value was reduced by RSD, from 430 to 371 \u0026micro;S cm\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e in DST and 1256 to 387 \u0026micro;S cm\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e in DSC. The concentrations of SOC, TN and DOC were 1.24\u0026minus;1.42, 1.05\u0026minus;1.09 and 1.09\u0026minus;4.70 times higher in RSD treatments compared to CK, respectively. The C/N ratio also significantly increased after RSD. There were no significant differences in soil TP and TK concentrations between RSD treatments and CK. After RSD, soil NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e concentration decreased from 83 to 7 mg N kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e soil in DST and 973 to 3 mg N kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e soil in DSC, while soil NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e concentration increased by 69- and 27- fold in DST and DSC, respectively. Soil AP and AK concentrations were significantly higher in RSD compared to CK, except for AP concentration in DST where no significant difference between RSD and CK was observed. The bacteria, AOA, AOB, \u003cem\u003enirK\u003c/em\u003e, \u003cem\u003enirS\u003c/em\u003e and \u003cem\u003enosZ\u003c/em\u003e abundances were significantly higher in RSD than CK, except in DST where there was no difference in \u003cem\u003enirK\u003c/em\u003e abundance between RSD and CK (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The fungi abundance in RSD was significantly lower compared to CK in DST, while in DSC it was not different from CK. The \u003cem\u003eF. oxysporum\u003c/em\u003e, \u003cem\u003eF. solani\u003c/em\u003e and \u003cem\u003eRalstonia solanacearum\u003c/em\u003e abundances were reduced significantly by RSD, except in DSC with no difference in \u003cem\u003eRalstonia solanacearum\u003c/em\u003e abundance between RSD and CK.\u003c/p\u003e \u003cp\u003eAfter planting, soil pH generally decreased (i.e CC\u0026thinsp;\u0026gt;\u0026thinsp;CP and RC\u0026thinsp;\u0026gt;\u0026thinsp;RP; Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e1\u003c/span\u003ea). The soil pH was significantly higher in RSD than CK, irrespectively of the presence of plants, i.e RC\u0026thinsp;\u0026gt;\u0026thinsp;CC and RP\u0026thinsp;\u0026gt;\u0026thinsp;CP. Plants significantly reduced soil DOC concentration in DST, while enhanced it in DSC (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e1\u003c/span\u003eb). Soil DOC concentration was significantly higher in CC than RC, while it was slightly higher in RP than CP but it was not significantly different in the two studied soils. After planting, no significant changes in the bacteria and AOA abundances were recorded (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e1\u003c/span\u003ec and e). RSD significantly enhanced the bacteria and AOA abundances (i.e. RC\u0026thinsp;\u0026gt;\u0026thinsp;CC and RP\u0026thinsp;\u0026gt;\u0026thinsp;CP). Bacterial abundance showed a positive correlation with soil pH (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ea). In general, the presence of plants enhanced fungal abundance (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e1\u003c/span\u003ed). The fungal abundance in RSD was significantly lower compared to CK in DST (i.e. RC\u0026thinsp;\u0026lt;\u0026thinsp;CC and RP\u0026thinsp;\u0026lt;\u0026thinsp;CP), whereas the opposite occurred in DSC (i.e. RC\u0026thinsp;\u0026gt;\u0026thinsp;CC and RP\u0026thinsp;\u0026gt;\u0026thinsp;CP). The AOB abundance was significantly higher in CC than CP, and RC than RP (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e1\u003c/span\u003ef). RSD significantly enhanced the AOB abundance regardless of the presence of plants, i.e RC\u0026thinsp;\u0026gt;\u0026thinsp;CC and RP\u0026thinsp;\u0026gt;\u0026thinsp;CP. In DST, the vegetable biomass, net photosynthetic rate and N uptake were slightly higher in RP than CP, but not significantly (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e1\u003c/span\u003eg-i). In DSC, vegetable biomass, net photosynthetic rate and N uptake in RP were1.33 g pot\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, 17.09 \u0026micro;mol m\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e s\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, and 46.93 mg N pot\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, respectively, and significantly higher than in CP (0.66 g pot\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, 6.29 \u0026micro;mol m\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e s\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, and 26.78 mg N pot\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eGross and net N transformation rates\u003c/h2\u003e \u003cp\u003eThe observed and modelled concentrations and \u003csup\u003e15\u003c/sup\u003eN abundances of NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e, NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e, and plant N matched well (R\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;\u0026gt;\u0026thinsp;0.94, Fig. S2-3). The \u003csup\u003e15\u003c/sup\u003eN- tracing analysis showed that soil gross N mineralization rates (\u003cem\u003eM\u003c/em\u003e) were 1.4\u0026minus;3.9 times higher in RSD (RC and RP) than CK (CC and CP; Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e2\u003c/span\u003ea). In DST, the presence of vegetables significantly reduced soil \u003cem\u003eM\u003c/em\u003e (i.e. CP\u0026thinsp;\u0026lt;\u0026thinsp;CC and RP\u0026thinsp;\u0026lt;\u0026thinsp;CP). There was no difference in soil \u003cem\u003eM\u003c/em\u003e between the presence and absence of vegetables in DSC, i.e CC \u0026asymp; CP and RC \u0026asymp; RP. Soil autotrophic nitrification rate (\u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e) was significantly lower in CC than RC (3.89 vs. 7.95 mg N kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e d\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e in DST and 10.30 vs. 20.09 mg N kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e d\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e in DSC), and CP than RP (0.85 vs. 3.61 mg N kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e d\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e in DST and 1.85 vs. 3.15 mg N kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e d\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e in DSC; Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e2\u003c/span\u003eb). The presence of vegetables significantly reduced soil \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e (i.e. CC\u0026thinsp;\u0026gt;\u0026thinsp;CP and RC\u0026thinsp;\u0026gt;\u0026thinsp;RP). Soil heterotrophic nitrification rate (\u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNrec\u003c/em\u003e\u003c/sub\u003e) were negligible in the absence of vegetables (i.e CC and RC), and increased to 0.81\u0026minus;2.61 mg N kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e d\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e in the presence of vegetables (i.e CP and RP). There were no differences in \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNrec\u003c/em\u003e\u003c/sub\u003e between CC and RC, and between CP and RP in both soils, except in DSC where the \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNrec\u003c/em\u003e\u003c/sub\u003e in RP was significantly higher than CP. \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e accounted for 30\u0026ndash;100% of soil gross N nitrification rates (\u003cem\u003eN\u003c/em\u003e, \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e + \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNrec\u003c/em\u003e\u003c/sub\u003e).\u003c/p\u003e \u003cp\u003eMicrobial NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e (\u003cem\u003eI\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e) and NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e (\u003cem\u003eI\u003c/em\u003e\u003csub\u003e\u003cem\u003eNO3\u003c/em\u003e\u003c/sub\u003e) immobilization rates (\u003cem\u003eI\u003c/em\u003e\u003csub\u003e\u003cem\u003eTN\u003c/em\u003e\u003c/sub\u003e) were negligible in the absence of vegetables (i.e CC and RC), while they significantly increased on average to 4.52 mg N kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e d\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e in the presence of vegetables (i.e CP and RP; Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e2\u003c/span\u003ec). No significant differences in \u003cem\u003eI\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e and \u003cem\u003eI\u003c/em\u003e\u003csub\u003e\u003cem\u003eNO3\u003c/em\u003e\u003c/sub\u003e were observed between RSD (RC) and CK (CC) in the absence of vegetables. In the presence of vegetables, the \u003cem\u003eI\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e and \u003cem\u003eI\u003c/em\u003e\u003csub\u003e\u003cem\u003eNO3\u003c/em\u003e\u003c/sub\u003e in RSD (i.e. RP, on average 4.13 mg N kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e d\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) were significantly lower than CK (i.e. CP, on average 0.62 mg N kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e d\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e). Plant NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e uptake rate (\u003cem\u003eU\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e) in RSD (RP) was 2.29 and 13.49 mg N kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e d\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e in DST and DSC, respectively, and significantly higher than in CK (CP, 1.59 and 2.06 mg N kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e d\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e in DST and DSC, respectively; Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e2\u003c/span\u003ed). Plant NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e uptake rate (\u003cem\u003eU\u003c/em\u003e\u003csub\u003e\u003cem\u003eNO3\u003c/em\u003e\u003c/sub\u003e) in RSD (RP) was not different from CK (CP) in DST, while it was significantly higher compared to CK in DSC (14.49 vs. 15.82 mg N kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e d\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e). Soil net N mineralization (\u003cem\u003eNet M\u003c/em\u003e) and nitrification (\u003cem\u003eNet N\u003c/em\u003e) rates was 1.8\u0026ndash;3.3 and 1.9\u0026ndash;2.0 times higher, respectively, in RC than CC (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e2\u003c/span\u003ee and f).\u003c/p\u003e \u003cp\u003eThe (\u003cem\u003eU\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e + \u003cem\u003eI\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e)/\u003cem\u003eM\u003c/em\u003e in RP was 9.2 and 1.1 folds lower in DST and DSC, respectively, compared to CP (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e2\u003c/span\u003eg). The presence of vegetables significantly enhanced the (\u003cem\u003eU\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e + \u003cem\u003eI\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e)/\u003cem\u003eM\u003c/em\u003e (i.e. CC\u0026thinsp;\u0026lt;\u0026thinsp;CP and RC\u0026thinsp;\u0026lt;\u0026thinsp;RP). The (\u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e + \u003cem\u003eI\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e)/\u003cem\u003eM\u003c/em\u003e was significantly lower in RC than CC, and RP than CP in both soils, except in DSC where no significant difference in this ratio between CC and RC was observed (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e2\u003c/span\u003eh). The presence of plants significantly enhanced the (\u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e + \u003cem\u003eI\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e)/\u003cem\u003eM\u003c/em\u003e in DST (i.e. CC\u0026thinsp;\u0026lt;\u0026thinsp;CP, RC\u0026thinsp;\u0026lt;\u0026thinsp;RP), and generally reduced this ratio in DSC. The NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e retention time ranged from 10.4 to 39.3 d in RC and RP, which was significantly higher than that in CC and CP, 4.6\u0026minus;13.1d; Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e2\u003c/span\u003ei). NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e retention time significantly higher in the presence of vegetables (i.e. CP and RP) than the absence of vegetables (i.e. CC and RC) in studied soils, except in DSC where the NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e retention time was significantly lower in CP than CC.\u003c/p\u003e \u003cp\u003e \u003cem\u003eM\u003c/em\u003e was significantly and positively correlated with the bacterial abundance (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05; Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eb) and soil pH (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.01; Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ec). \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e increased significantly with increasing soil pH (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.01; Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ed), and \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e showed a significantly negative correction with the ratio of (\u003cem\u003eU\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e + \u003cem\u003eI\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e)/\u003cem\u003eM\u003c/em\u003e (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.01; Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ee). There was a significantly positive relationship between \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNrec\u003c/em\u003e\u003c/sub\u003e and fungal abundance (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05; Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ef).\u003c/p\u003e \u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003eThe results of the present study show that RSD significantly alters the gross N transformations and increases plant N uptake rates of soils that were used for continuous vegetable production. Consistent with our hypothesis, RSD significantly stimulated soil gross N mineralization and autotrophic nitrification rates, which enhanced the supply of N of both degraded soils and subsequent N uptake by plants. RSD improved soil quality (i.e. increasing soil pH, and decreasing EC and NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e accumulation), further promoting vegetable N uptake, being in line with our hypotheses.\u003c/p\u003e \u003cdiv id=\"Sec10\" class=\"Section2\"\u003e \u003ch2\u003eRSD accelerated the production of soil NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e\u003c/h2\u003e \u003cp\u003eOrganic N mineralization to NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e and subsequent oxidation to NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e (autotrophic nitrification, \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e) as well as direct oxidation of organic N to NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e (heterotrophic nitrification, \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNrec\u003c/em\u003e\u003c/sub\u003e) were the two important pathways for NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e production (M\u0026uuml;ller et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Zhang et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). In this study, soil \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e was the major contributor producing NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e, particularly in the absence of vegetables (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e2\u003c/span\u003eb). We found that RSD significantly enhanced \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e irrespectively of plants. Generally, soil N mineralization dominates NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e production and is a key factor affecting \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e (Elrys et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2021b\u003c/span\u003e; Schimel and Bennett \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). In this study, Soil gross N mineralization (\u003cem\u003eM\u003c/em\u003e) was significantly higher in RSD compared to CK and affected by plants (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e2\u003c/span\u003ea), indicating that RSD did stimulate soil \u003cem\u003eM\u003c/em\u003e and subsequent NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e production. This could be a main reason leading to higher soil NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e concentration in RSD than CK (Table\u0026nbsp;1). It is widely reported that RSD can alter soil properties (Huang et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Zhu et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2014\u003c/span\u003e), which is consistent with our results where soil pH, the concentrations of SOC, TN and DOC, and the bacteria abundance were enhanced by RSD (Table\u0026nbsp;1; Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e1\u003c/span\u003ea, b and c). Numerous studies have shown that above-mentioned indicators were key for the regulation of \u003cem\u003eM\u003c/em\u003e, with often significantly positive relationships (Elrys et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2021a\u003c/span\u003e; Wang et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Zhang et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). We confirmed this by a positive correlation of \u003cem\u003eM\u003c/em\u003e with soil pH and the bacterial abundance (Fig.\u0026nbsp;4b and c). Furthermore, higher soil DOC, SOC and TN concentrations stimulated by RSD which in turn also affected soil pH and bacterial abundance led to increased \u003cem\u003eM\u003c/em\u003e and therefore higher NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e production. This is in line with the general observation that a higher NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e production via \u003cem\u003eM\u003c/em\u003e is also enhancing \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e, explaining the close correlation between \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e and \u003cem\u003eM\u003c/em\u003e (Elrys et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2021b\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn addition, NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e is also the N sources of vegetables and microorganisms in vegetable-soil systems (Hodge et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Inselsbacher et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Kuzyakov and Xu \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Therefore, vegetable NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e uptake (\u003cem\u003eU\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e) and microbial NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e immobilization (\u003cem\u003eI\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e) also affect \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e. Lower (\u003cem\u003eU\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e + \u003cem\u003eI\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e)/\u003cem\u003eM\u003c/em\u003e ratio in RSD compared to CK can be caused by lower \u003cem\u003eU\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e and \u003cem\u003eI\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e or higher \u003cem\u003eM\u003c/em\u003e, both enhancing the availability of NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e to ammonia-oxidizing microorganisms which in turn stimulates \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e. This was supported by significantly negative relationships between \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e and (\u003cem\u003eU\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e + \u003cem\u003eI\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e)/\u003cem\u003eM\u003c/em\u003e in this study (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ee). Soil pH is a key factor regulating \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e (Elrys et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2021b\u003c/span\u003e; Shammas \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e1986\u003c/span\u003e) and a significantly positive relationship between \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e and soil pH existed in this study (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ec). Increasing soil pH can promote the release of NH\u003csub\u003e3\u003c/sub\u003e and enhance the activities of ammonia-oxidizing microorganisms (e.g. AOA and AOB), which has been shown to stimulate \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e (Li et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Norton and Stark \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Stempfhuber et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Here we provided evidence that soil pH and the AOA and AOB abundances were enhanced by RSD and in turn increased \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e.\u003c/p\u003e \u003cp\u003eWe also confirmed a stimulation of \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNrec\u003c/em\u003e\u003c/sub\u003e by RSD as shown by Zhu et al. (\u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2014\u003c/span\u003e), but only when plants were present. The exudates (e.g. acetic acids associated with \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNrec\u003c/em\u003e\u003c/sub\u003e) can be metabolized instantaneously and decomposed by 40% within two days (Laughlin et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2008\u003c/span\u003e) explaining for instance why \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNrec\u003c/em\u003e\u003c/sub\u003e decreased from 0.34 to 0 mg N kg d\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e at room temperature (He et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) and can explain negligible \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNrec\u003c/em\u003e\u003c/sub\u003e in the absence of vegetables after 26 and 18 days cultivation. Surprisingly, RSD significantly enhanced \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNrec\u003c/em\u003e\u003c/sub\u003e in the presence of vegetables (especially in DSC). After RSD treatment, soil NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e content was reduced, and the NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e production via \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e (ranging from 3.15 to 3.61 mg N kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e d\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) was not sufficient to meet plant NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e uptake rates (ranging from 8.59 to 15.82 mg N kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e d\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e). Thus, stimulation of \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNrec\u003c/em\u003e\u003c/sub\u003e is one way to meet the N demand of the plants. These results indicate that \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNrec\u003c/em\u003e\u003c/sub\u003e can be an important pathway for N supply to plants, which was in line with previous studies (Dan et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; He et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAlthough both vegetables and microorganisms can assimilate soil NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e, increasing soil NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e production rates (\u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e + \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNrec\u003c/em\u003e\u003c/sub\u003e) due to RSD resulted in higher soil net nitrification rates compared to CK (positive relationship, Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ef), particularly in the absence of vegetables, which increased the leaching and accumulation risks of NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e. This indicated that soil N fertilizer management after RSD treatment need to be carefully managed to avoid NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e accumulation.\u003c/p\u003e \u003cp\u003e \u003cem\u003eRSD promoted vegetable N uptake by regulating soil gross N transformation and improving quality of degraded vegetable soils\u003c/em\u003e \u003c/p\u003e \u003cp\u003eIn this study, vegetable NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e (\u003cem\u003eU\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e) and NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e (\u003cem\u003eU\u003c/em\u003e\u003csub\u003e\u003cem\u003eNO3\u003c/em\u003e\u003c/sub\u003e) uptake rates were generally higher in RP than CP (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e2\u003c/span\u003ed), indicating that RSD did promote vegetable N uptake. We found that significant differences in vegetable N uptake rates (\u003cem\u003eU\u003c/em\u003e\u003csub\u003e\u003cem\u003eTN\u003c/em\u003e\u003c/sub\u003e, \u003cem\u003eU\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e + \u003cem\u003eU\u003c/em\u003e\u003csub\u003e\u003cem\u003eNO3\u003c/em\u003e\u003c/sub\u003e) between RP and CP were mainly attributed to differences in \u003cem\u003eU\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e. Although studied vegetables (i.e. tomato and cucumber) are characterized by NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e-N preference, NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e-N assimilation for vegetables is not only more energy efficient than NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e-N, but also can promote root proliferation and keep the ionic balance (Al-Harbi \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Bloom et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; George et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Nacry et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Soil gross N mineralization (\u003cem\u003eM\u003c/em\u003e), autotrophic nitrification (\u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e) and NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e immobilization (\u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e) are key processes to govern soil NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e concentration and NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e retention time (Schimel and Bennett \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2004\u003c/span\u003e), thereby affecting \u003cem\u003eU\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e. A lower (\u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e \u003cem\u003e+ I\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e)/\u003cem\u003eM\u003c/em\u003e ratio in RP than CP causes a higher NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e production (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e2\u003c/span\u003eh), leading to increased soil NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e concentrations and in turn can promote \u003cem\u003eU\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e. In addition, compared to CP, longer NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e retention time due higher soil initial NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e concentration or lower \u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e and \u003cem\u003eI\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e in RP (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e2\u003c/span\u003eg) can be an important factor responsible for higher \u003cem\u003eU\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e. Despite lower soil NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e concentration in RSD compared to CK, \u003cem\u003eU\u003c/em\u003e\u003csub\u003e\u003cem\u003eNO3\u003c/em\u003e\u003c/sub\u003e was generally higher in RP than CP (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e2\u003c/span\u003ed), which was likely duo to increasing soil gross nitrification rates after RSD. Higher soil gross nitrification rates in RP than CP (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e2\u003c/span\u003eb) can produce more NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e for plant N uptake, explaining a stimulation of \u003cem\u003eU\u003c/em\u003e\u003csub\u003e\u003cem\u003eNO3\u003c/em\u003e\u003c/sub\u003e. Clearly, RSD promoted vegetable N uptake by regulating soil gross N transformation processes in this study.\u003c/p\u003e \u003cp\u003eIn addition to soil N transformations, RSD can also alter soil physiochemical properties and thus influence the N uptake of plants. Soil physiochemical properties (e.g. soil pH, EC, microorganisms and structure) are important factors affecting vegetable growth (Bernstein \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e1975\u003c/span\u003e; Lau and Lennon \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Neina \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Passioura \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e1991\u003c/span\u003e). Long-term continuous cropping of the same or similar crop variety and high N fertilizer input can degrade the cultivated soil, e.g. soil NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e accumulation, acidification and salinization, and increase of the abundances and diversities of soil pathogens, which can seriously affect plant growth (Guo et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Hu et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Huang et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Shipton \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e1977\u003c/span\u003e). In this study, the quality of degraded soil was improved by RSD, i.e. increasing of soil pH and decreasing the contents of soil EC, NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e and the abundances of pathogens (\u003cem\u003eF. oxysporum\u003c/em\u003e, \u003cem\u003eF. solani\u003c/em\u003e and \u003cem\u003eRalstonia solanacearum\u003c/em\u003e; Table\u0026nbsp;1), which supports plant growth, a higher net photosynthetic rate, total N uptake (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e1\u003c/span\u003eg-i and \u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e2\u003c/span\u003ed).\u003c/p\u003e \u003c/div\u003e"},{"header":"Conclusions","content":"\u003cp\u003eWe were able to show that RSD significantly promotes plant N uptake, which can be attributed to two properties: 1) RSD decreased (\u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e \u003cem\u003e+ I\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e)/\u003cem\u003eM\u003c/em\u003e, and increased NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e retention time and production rates of soil NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e, enhancing the N supply capacity of degraded soils; 2) RSD improved quality of degraded soil (i.e. increasing soil pH and decreasing the contents of soil EC and NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e and pathogen abundances). Increasing soil net nitrification rates indicated that RSD accelerated soil NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e production in the soil and increased the risk of leaching or accumulation of NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e. Although RSD can improve the quality of degraded soils and promote plant N uptake, N fertilizer management after RSD still needs attention, otherwise NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e accumulation or leaching will occur more rapidly.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgements\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis work was supported by the National Natural Science Foundation of China (grant number 41830642),\u0026nbsp;the\u0026nbsp;Postgraduate Research and Practice Innovation Program of Jiangsu Province (KYCX21_1332),\u0026nbsp;and the “Double World-Classes”: Development\u0026nbsp;in\u0026nbsp;Geography Project. The study was carried out as part of the IAEA funded\u0026nbsp;coordinated\u0026nbsp;research\u0026nbsp;project “Minimizing farming impacts on climate change by enhancing carbon and nitrogen capture and storage in Agro-Ecosystems (D1.50.16)” and was carried out in close collaboration with the German Science\u0026nbsp;Foundation research unit DASIM (FOR 2337).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting interests:\u003c/strong\u003e None\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAkhtar M (2000) Roles of organic soil amendments and soil organisms in the biological control of plant-parasitic nematodes: a review. 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Plant Soil 382:269\u0026ndash;280. doi: \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s11104-014-2160-3\u003c/span\u003e\u003cspan address=\"10.1007/s11104-014-2160-3\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"},{"header":"Tables","content":"\u003cp\u003e\u003cstrong\u003eTable 1.\u003c/strong\u003e Soil chemical properties in different treatments.\u003c/p\u003e\n\u003ctable border=\"1\" cellpadding=\"0\" cellspacing=\"0\" width=\"0\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" width=\"4.644268774703558%\"\u003e\n \u003cp\u003eSoils\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"7.41106719367589%\"\u003e\n \u003cp\u003etreatments\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.399209486166008%\"\u003e\n \u003cp\u003epH\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"6.521739130434782%\"\u003e\n \u003cp\u003eEC\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.49802371541502%\"\u003e\n \u003cp\u003eSOC\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.399209486166008%\"\u003e\n \u003cp\u003eTN\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.399209486166008%\"\u003e\n \u003cp\u003eC/N\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.399209486166008%\"\u003e\n \u003cp\u003eTP\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"5.632411067193676%\"\u003e\n \u003cp\u003eTK\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"7.509881422924901%\"\u003e\n \u003cp\u003eDOC\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"5.632411067193676%\"\u003e\n \u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"6.521739130434782%\"\u003e\n \u003cp\u003eNO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e-\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"1.5810276679841897%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"5.928853754940712%\"\u003e\n \u003cp\u003eAP\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"6.521739130434782%\"\u003e\n \u003cp\u003eAK\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" width=\"4.644268774703558%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"7.41106719367589%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.399209486166008%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"6.521739130434782%\"\u003e\n \u003cp\u003e\u0026micro;S cm\u003csup\u003e-1\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" valign=\"top\" width=\"16.897233201581027%\"\u003e\n \u003cp\u003eg kg\u003csup\u003e-1\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.399209486166008%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" valign=\"top\" width=\"14.031620553359684%\"\u003e\n \u003cp\u003eg kg\u003csup\u003e-1\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"7.509881422924901%\"\u003e\n \u003cp\u003emg kg\u003csup\u003e-1\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" valign=\"top\" width=\"12.154150197628459%\"\u003e\n \u003cp\u003emg N kg\u003csup\u003e-1\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"1.5810276679841897%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" valign=\"top\" width=\"12.450592885375494%\"\u003e\n \u003cp\u003emg kg\u003csup\u003e-1\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" width=\"4.644268774703558%\"\u003e\n \u003cp\u003eDST\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"7.41106719367589%\"\u003e\n \u003cp\u003eCK\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.399209486166008%\"\u003e\n \u003cp\u003e5.81\u0026plusmn;0.03b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"6.521739130434782%\"\u003e\n \u003cp\u003e430\u0026plusmn;6a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.49802371541502%\"\u003e\n \u003cp\u003e12.19\u0026plusmn;0.12b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.399209486166008%\"\u003e\n \u003cp\u003e1.61\u0026plusmn;0.06b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.399209486166008%\"\u003e\n \u003cp\u003e7.56\u0026plusmn;0.29b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.399209486166008%\"\u003e\n \u003cp\u003e1.40\u0026plusmn;0.06a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"5.632411067193676%\"\u003e\n \u003cp\u003e19\u0026plusmn;0a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"7.509881422924901%\"\u003e\n \u003cp\u003e151\u0026plusmn;3b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"5.632411067193676%\"\u003e\n \u003cp\u003e1\u0026plusmn;0b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"6.521739130434782%\"\u003e\n \u003cp\u003e83\u0026plusmn;1a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"1.5810276679841897%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"5.928853754940712%\"\u003e\n \u003cp\u003e191\u0026plusmn;3a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"6.521739130434782%\"\u003e\n \u003cp\u003e260\u0026plusmn;4b\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" width=\"4.644268774703558%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"7.41106719367589%\"\u003e\n \u003cp\u003eRSD\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.399209486166008%\"\u003e\n \u003cp\u003e7.27\u0026plusmn;0.00a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"6.521739130434782%\"\u003e\n \u003cp\u003e371\u0026plusmn;3b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.49802371541502%\"\u003e\n \u003cp\u003e15.16\u0026plusmn;0.12a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.399209486166008%\"\u003e\n \u003cp\u003e1.76\u0026plusmn;0.03a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.399209486166008%\"\u003e\n \u003cp\u003e8.60\u0026plusmn;0.15a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.399209486166008%\"\u003e\n \u003cp\u003e1.41\u0026plusmn;0.12a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"5.632411067193676%\"\u003e\n \u003cp\u003e19\u0026plusmn;1a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"7.509881422924901%\"\u003e\n \u003cp\u003e164\u0026plusmn;6a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"5.632411067193676%\"\u003e\n \u003cp\u003e69\u0026plusmn;1a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"6.521739130434782%\"\u003e\n \u003cp\u003e7\u0026plusmn;0b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"1.5810276679841897%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"5.928853754940712%\"\u003e\n \u003cp\u003e190\u0026plusmn;5a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"6.521739130434782%\"\u003e\n \u003cp\u003e547\u0026plusmn;4a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" width=\"4.644268774703558%\"\u003e\n \u003cp\u003eDSC\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"7.41106719367589%\"\u003e\n \u003cp\u003eCK\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.399209486166008%\"\u003e\n \u003cp\u003e7.49\u0026plusmn;0.01b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"6.521739130434782%\"\u003e\n \u003cp\u003e1256\u0026plusmn;5a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.49802371541502%\"\u003e\n \u003cp\u003e15.12\u0026plusmn;0.47b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.399209486166008%\"\u003e\n \u003cp\u003e2.59\u0026plusmn;0.02b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.399209486166008%\"\u003e\n \u003cp\u003e5.83\u0026plusmn;0.18b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.399209486166008%\"\u003e\n \u003cp\u003e2.38\u0026plusmn;0.02a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"5.632411067193676%\"\u003e\n \u003cp\u003e22\u0026plusmn;0a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"7.509881422924901%\"\u003e\n \u003cp\u003e57\u0026plusmn;3b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"5.632411067193676%\"\u003e\n \u003cp\u003e6\u0026plusmn;08b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"6.521739130434782%\"\u003e\n \u003cp\u003e973\u0026plusmn;11a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"1.5810276679841897%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"5.928853754940712%\"\u003e\n \u003cp\u003e97\u0026plusmn;4b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"6.521739130434782%\"\u003e\n \u003cp\u003e131\u0026plusmn;0b\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" width=\"4.644268774703558%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"7.41106719367589%\"\u003e\n \u003cp\u003eRSD\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.399209486166008%\"\u003e\n \u003cp\u003e8.50\u0026plusmn;0.00a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"6.521739130434782%\"\u003e\n \u003cp\u003e387\u0026plusmn;16b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.49802371541502%\"\u003e\n \u003cp\u003e21.52\u0026plusmn;0.36a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.399209486166008%\"\u003e\n \u003cp\u003e2.71\u0026plusmn;0.03a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.399209486166008%\"\u003e\n \u003cp\u003e7.94\u0026plusmn;0.13a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.399209486166008%\"\u003e\n \u003cp\u003e2.38\u0026plusmn;0.05a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"5.632411067193676%\"\u003e\n \u003cp\u003e23\u0026plusmn;1a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"7.509881422924901%\"\u003e\n \u003cp\u003e268\u0026plusmn;16a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"5.632411067193676%\"\u003e\n \u003cp\u003e159\u0026plusmn;5a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"6.521739130434782%\"\u003e\n \u003cp\u003e3\u0026plusmn;0b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"1.5810276679841897%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"5.928853754940712%\"\u003e\n \u003cp\u003e203\u0026plusmn;9a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"6.521739130434782%\"\u003e\n \u003cp\u003e507\u0026plusmn;6a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003eDST, degraded tomato soil under tomato continuous cropping; DSC, degraded cucumber soil under cucumber continuous cropping; CK, a control (without reductive soil disinfestation); RSD, reductive soil disinfestation. EC, electrical conductivity; SOC, soil organic carbon; TN, total nitrogen; C/N, SOC: TN ratio; TP, total phosphorus; TK, total potassium; NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e,\u0026nbsp;ammonium;\u0026nbsp;NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026ndash;\u003c/sup\u003e,\u0026nbsp;nitrate;\u0026nbsp;AP,\u0026nbsp;available phosphorus; AK,\u0026nbsp;available\u0026nbsp;potassium.\u0026nbsp;The different lowercase letters indicate the differences between CK and RSD in the same degraded soil (\u003cem\u003ep\u003c/em\u003e \u0026lt;0.05).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTable 2.\u003c/strong\u003e Soil microbial properties in different treatments.\u003c/p\u003e\n\u003ctable border=\"1\" cellpadding=\"0\" cellspacing=\"0\" width=\"0\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" width=\"4.577114427860696%\"\u003e\n \u003cp\u003eSoils\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.059701492537313%\"\u003e\n \u003cp\u003etreatments\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"9.45273631840796%\"\u003e\n \u003cp\u003eBacteria\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003eFungi\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003eAOA\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003eAOB\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003enirK\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003enirS\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003enosZ\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003eF.o\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003eF.s\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003eR.s\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" width=\"4.568023833167826%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.04369414101291%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"10\" valign=\"top\" width=\"87.38828202581927%\"\u003e\n \u003cp\u003elog\u003csub\u003e10\u003c/sub\u003e gene copies g\u003csup\u003e-1\u003c/sup\u003e dry soil\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" width=\"4.577114427860696%\"\u003e\n \u003cp\u003eDST\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.059701492537313%\"\u003e\n \u003cp\u003eCK\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"9.45273631840796%\"\u003e\n \u003cp\u003e9.88\u0026plusmn;0.04b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e9.03\u0026plusmn;0.01a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e7.00\u0026plusmn;0.05b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e6.96\u0026plusmn;0.04b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e7.86\u0026plusmn;0.16a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e7.30\u0026plusmn;0.04b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e7.92\u0026plusmn;0.04b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e5.14\u0026plusmn;0.14a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e6.17\u0026plusmn;0.09a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e6.03\u0026plusmn;0.15a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" width=\"4.577114427860696%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.059701492537313%\"\u003e\n \u003cp\u003eRSD\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"9.45273631840796%\"\u003e\n \u003cp\u003e10.39\u0026plusmn;0.03a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e8.75\u0026plusmn;0.05b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e7.85\u0026plusmn;0.03a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e7.97\u0026plusmn;0.03a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e7.89\u0026plusmn;0.11a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e8.33\u0026plusmn;0.02a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e9.20\u0026plusmn;0.04a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e4.48\u0026plusmn;0.13b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e4.61\u0026plusmn;0.05b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e5.48\u0026plusmn;0.13b\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" width=\"4.577114427860696%\"\u003e\n \u003cp\u003eDSC\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.059701492537313%\"\u003e\n \u003cp\u003eCK\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"9.45273631840796%\"\u003e\n \u003cp\u003e10.17\u0026plusmn;0.06b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e8.29\u0026plusmn;0.08a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e8.15\u0026plusmn;0.03b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e6.52\u0026plusmn;0.10b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e7.01\u0026plusmn;0.12b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e7.38\u0026plusmn;0.08b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e7.52\u0026plusmn;0.07b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e5.14\u0026plusmn;0.19a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e4.67\u0026plusmn;0.10a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e5.72\u0026plusmn;0.19a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" width=\"4.577114427860696%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.059701492537313%\"\u003e\n \u003cp\u003eRSD\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"9.45273631840796%\"\u003e\n \u003cp\u003e10.85\u0026plusmn;0.05a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e8.31\u0026plusmn;0.04a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e8.35\u0026plusmn;0.01a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e6.98\u0026plusmn;0.05a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e7.84\u0026plusmn;0.15a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e8.24\u0026plusmn;0.09a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e8.64\u0026plusmn;0.10a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e4.81\u0026plusmn;0.05b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e4.11\u0026plusmn;0.09b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"8.656716417910447%\"\u003e\n \u003cp\u003e5.79\u0026plusmn;0.07a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003eDST, degraded tomato soil under tomato continuous cropping; DSC, degraded cucumber soil under cucumber continuous cropping; CK, a control (without reductive soil disinfestation); RSD, reductive soil disinfestation. AOA, ammonia-oxidizing archaea; AOB, ammonia-oxidizing bacteria; \u003cem\u003enirK\u003c/em\u003e and \u003cem\u003enirS\u003c/em\u003e, the genes of nitrite reductase; \u003cem\u003enosZ\u003c/em\u003e, the gene of nitrous oxide reductase.\u0026nbsp;The different lowercase letters indicate the differences between CK and RSD in the same degraded vegetable soil (\u003cem\u003ep\u003c/em\u003e \u0026lt;0.05).\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":true,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"plant-and-soil","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"plso","sideBox":"Learn more about [Plant and Soil](https://www.springer.com/journal/11104)","snPcode":"11104","submissionUrl":"https://submission.nature.com/new-submission/11104/3","title":"Plant and Soil","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Reductive soil disinfestation, vegetable N uptake, Gross N transformation rates, degraded vegetable soils, 15N tracing","lastPublishedDoi":"10.21203/rs.3.rs-1805779/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-1805779/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003ch2\u003eAims\u003c/h2\u003e \u003cp\u003eReductive soil disinfestation (RSD) has been widely applied to improve soil degradation, thereby enhancing vegetable N uptake and subsequently productivity. However, the effect of RSD on interactions between vegetable N uptake and soil gross N transformation remain unclear.\u003c/p\u003e\u003ch2\u003eMethods\u003c/h2\u003e \u003cp\u003eTwo degraded vegetable soils were treated with RSD. Untreated soils served as control (CK). To quantify the effects of RSD on N cycling in vegetable-soil systems, \u003csup\u003e15\u003c/sup\u003eN tracing pot experiments were conducted.\u003c/p\u003e\u003ch2\u003eResults\u003c/h2\u003e \u003cp\u003eVegetable NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e and NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e uptake rates were 1.0\u0026ndash;6.5 times higher in RSD treatments than CK. Soil gross N mineralization rates (\u003cem\u003eM\u003c/em\u003e) in RSD ranged from 0.83 to 13.00 mg N kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e d\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, and were significantly higher than in CK (0.21 to 8.71 mg N kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e d\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e). Autotropic nitrification rates (\u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e) increased by 1.7\u0026ndash;4.2 times after RSD. NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e immobilization rates (\u003cem\u003eI\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e) were significantly inhibited by RSD in the presence of vegetables. These induced decreasing (\u003cem\u003eO\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e \u003cem\u003e+ I\u003c/em\u003e\u003csub\u003e\u003cem\u003eNH4\u003c/em\u003e\u003c/sub\u003e)/\u003cem\u003eM\u003c/em\u003e, increasing NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e retention times and production rates of soil NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e after RSD treatment. Thus, RSD promoted N supply to vegetables and subsequently N uptake by vegetables on degraded soils. In addition, RSD improved the quality on degraded soils (i.e. increasing soil pH, and decreasing soil EC and NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e contents and pathogen abundances), which could also be an important factor promoting vegetable N uptake.\u003c/p\u003e\u003ch2\u003eConclusion\u003c/h2\u003e \u003cp\u003eThus, RSD can promote vegetable N uptake by regulating soil gross N transformations and improving the quality of degraded vegetable soils. However, N fertilizer management still needs attention because of stimulated NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e production rates after RSD, which may lead to more rapid NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e leaching or gaseous N losses.\u003c/p\u003e","manuscriptTitle":"Reductive soil disinfestation promoted vegetable N uptake by regulating soil gross N transformations and improving the quality of degraded vegetable soils","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2022-07-12 16:36:19","doi":"10.21203/rs.3.rs-1805779/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Minor revisions","date":"2022-07-24T18:14:39+00:00","index":"","fulltext":""},{"type":"reviewerAgreed","content":"","date":"2022-07-05T00:37:13+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2022-07-04T21:21:31+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"Plant and Soil","date":"2022-07-01T06:12:02+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2022-07-01T06:10:06+00:00","index":"","fulltext":""},{"type":"submitted","content":"Plant and Soil","date":"2022-06-28T22:35:53+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"plant-and-soil","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"plso","sideBox":"Learn more about [Plant and Soil](https://www.springer.com/journal/11104)","snPcode":"11104","submissionUrl":"https://submission.nature.com/new-submission/11104/3","title":"Plant and Soil","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"f5ec3fdb-a573-4010-a809-15a56fa543f0","owner":[],"postedDate":"July 12th, 2022","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2022-08-30T08:59:43+00:00","versionOfRecord":[],"versionCreatedAt":"2022-07-12 16:36:19","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-1805779","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-1805779","identity":"rs-1805779","version":["v1"]},"buildId":"-HB7Z8yhvgn0wM9Nzuekk","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}