Phosphorus-arsenic interaction mitigates toxicity and accumulation of arsenic in rice grown in contaminated fields

Phosphorus-arsenic interaction mitigates toxicity and accumulation of arsenic in rice grown in contaminated fields | 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 Phosphorus-arsenic interaction mitigates toxicity and accumulation of arsenic in rice grown in contaminated fields Kavita Shukla, Saurabh Kumar Pathak, Swarnendra Banjerjee, Sudhakar Srivastava This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8891770/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 16 You are reading this latest preprint version Abstract Arsenic (As) contamination in rice is a serious threat to food security worldwide. Phosphorus (P) plays a crucial role in modulating As uptake in rice plants. Elevated P content in the soil may hinder the uptake of As by competing with the transporter system and altering rhizosphere chemistry. This study aimed to investigate the effect of varied P fertilizer levels on rice ( Oryza sativa L.) plants grown in As-contaminated soil in terms of growth and As accumulation. The experiment included recommended (60 kg h -1 ), half (30 kg h -1 ), and double (120 kg h -1 ) doses of diammonium phosphate (DAP). Two contrasting varieties of rice were used that differed in phosphorus use efficiency (PUE): P-efficient DNA Sribala (DS) and P-inefficient Sai Kasturi (SK). SK suffered significant reductions in shoot and panicle lengths under low P doses (up to 35% and 18%, respectively), whereas DS showed non-significant effects. Arsenic accumulation in grains increased by 20% under low P doses for SK but decreased by 9-11% with higher P doses. In contrast, DS demonstrated tolerance to low phosphorus concentrations and no significant change in As accumulation was observed. In addition, the yield of rice was also increased in high P doses, which was 14-23% at two sites for SK, while 17-20% for DS. These findings underscore the critical role of P management in mitigating As toxicity and optimizing rice productivity in contaminated environments. Antioxidant enzymes Crop productivity Food safety Phosphorus accumulation Phosphorus use efficiency Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 Introduction The presence of hazardous metal(loid) elements in soil and water is a growing global concern, posing serious threats to human and animal health and negatively impacting the well-being of plants and microorganisms (Majumdar et al., 2023; Tyagi et al., 2022 ). Unlike organic pollutants, metal(loid)s do not degrade and persist in soil for prolonged durations (Wang et al., 2021 ; Wang et al., 2022 ). Soil enriched with arsenic (As) is a significant source of contamination in the food chain and water supply, raising environmental and human health concerns due to the fact that As is a well-established carcinogen and mutagen (Chakrabarty 2015 ). Several soil parameters can influence As accumulation in rice, including total soil As concentration, redox status, poorly crystalline iron (Fe) (hydr)oxide concentration, plant available phosphorus (P), texture, pH, plant available silicon (Si), and sulfur (S) (Linam et al., 2024 ; Amaral et al., 2017 ). Phosphorus plays a critical role in regulating the uptake of As by plants through two primary processes. Arsenate [As(V)] serves as a phosphate analogue and is taken up by plants through a phosphate transporter system (Latowski et al., 2018 ). Therefore, As uptake and accumulation are known to be inversely influenced by P concentrations in the soil solution (Wu et al., 2022 ). Arsenic in soil exists in both inorganic and organic forms, including arsenite As(III), arsenate (As(V)), monomethylarsonic acid (MMA), and dimethylarsonic acid (DMA). Typically, As(V) is present as H 2 AsO 3 −3 or HAsO 2 −4 , and As(III) as H 3 AsO 3 under circumneutral pH conditions (Huang et al., 2024 ; Seyfferth et al., 2017 ). In the soil, phosphate competes for the same sorption sites as As(V) (Rivas-Pérez et al., 2015 ). This competition facilitates the desorption of As into the soil solution, enabling its uptake by plants (Bakhat et al., 2017; Rahman and Hesegawa, 2011). Therefore, the impact of P on As mobilization depends on the soil's charge characteristics. In a hydroponic experiment, As-induced reductions in root growth and increased tissue As concentration were reversed by P treatment (Tu and Ma, 2003 ). Therefore, P can effectively reduce As availability and root uptake in As-contaminated soils (Anwar et al., 2015). Cytoplasmic As(V) can disrupt metabolic processes involving phosphate, making it toxic to plants (Meharg and Hartley-Whitaker, 2002 ). However, when internal phosphate concentration is high, it can lead to the down-regulation of phosphate transporters and reduce As uptake (Finnegan and Chen, 2012 ). Although As is not a redox-active metal, substantial evidence suggests that exposing plants to inorganic As generates reactive oxygen species (ROS), connected with valence changes of As from As(V) to As(III) in plants (Abbas et al., 2018 ; Meharg and Hartley-Whitaker, 2002 ). The chemical nature of As(V) and phosphate, and their interactions in soil and plants, do indicate that P amendment can be a fruitful and practical strategy to regulate As levels in plants. However, it is important to exercise caution when amending soils with P to mitigate As uptake. In soils with low P concentrations and high As levels, increasing soil P levels can elevate As solubility, availability, uptake, and accumulation in rice grain due to P adsorption to soil exchange sites. Therefore, a balanced approach is required when using P to manage As contamination in the soil. In our previous work, two varieties differing in phosphorus use efficiency (PUE) were identified, and their interactions with As were studied in laboratory conditions (Shukla et al., 2024 ). Sai Kasturi (SK) was identified as a low PUE while DNA Sribala (DS) was identified as a high PUE variety. This study attempted to investigate phosphorus fertilization effects on arsenic uptake, oxidative stress responses, and crop productivity in rice crops under arsenic-contaminated soil conditions. Using two different varieties of rice defined by phosphorus utilization efficiency (PUE), research attempted to define genotype-related responses to varied phosphate supplies. The main goal was to find out if optimized phosphorus management, as a means to minimize arsenic content in rice grains, can still allow crop growth and production, providing an achievable and sustainable crop production practice to support food security in arsenic-affected areas. Material and Methods Experimental Site and Rice Cultivar Selection In the study, As-contaminated fields were chosen to examine the effects of different P doses on particular rice types. The study was conducted in two different districts of Uttar Pradesh, India: Ghazipur (site 1) and Jaunpur (site 2) (Supplementary Figure S1 ). These fields, which are located at 25.5084° N, 93.8925° E for Ghazipur and 25°34'22.1" N, 82°58'32.0" E for Jaunpur, were selected because of their comparable levels of contamination: Ghazipur (17.3 mg kg − 1 ) and Jaunpur (16.5 mg kg − 1 ). Two selected rice cultivars were DNA sribala (DS) and Sai kasturi (SK), which were identified in our earlier lab study (Shukla et al., 2024 ). DNA Sribala is a high PUE variety, responsive to phosphorus and tolerant to As stress under hydroponic conditions. Sai Kasturi is the low PUE variety and is As sensitive too (Shukla et al., 2024 ). These rice varieties typically take between 125 to 135 days to reach maturity. Field Layout, Irrigation, Experimental Design, and Fertilizer Application The entire experiment was executed using a completely randomized block design. The field sites at two locations were divided into three blocks. The control block (normal As-contaminated soil at two sites) received a P dose of 60 kg ha − 1 . There were two experimental blocks: experimental blocks P1 (As-contaminated soil amended with 30 kg ha − 1 of P; half of the recommended P fertilizer) and P2 (As-contaminated soil amended with 120 kg ha − 1 of P; double the recommended P fertilizer). The seeds of both varieties were grown in nursery where soil-As concentration was less than 5 mg kg − 1 . Two square shape (5*5 ft 2 ) plots were prepared with submerged water condition (2–3 cm). After 30 days after sowing (DAS) the rice plants with equal height were transferred into different experimental plots. Throughout the growth period (July to November), the control and experimental fields were kept at a constant water level of 3 to 6 cm. The ratio of (N:P: K) fertilizer applied per hectare was 120:60:40. (N:P: K) in control block, 120:30:40 kg ha − 1 in P1 and 120:120:40 kg ha − 1 in P2 with urea, superphosphate and muriate of potash being the respective sources. Nitrogen was applied in three stages: half during field preparation, and the remaining quarters were given during the early growth stage (tillering, 25DAS) and 50–60 days after transplant (DAT). Weeding was managed manually four times throughout the entire experimental period, with the waterlogged conditions also contributing to weed control in the fields. Soil Sampling and Analysis Arsenic-contaminated topsoil (0–20 cm depth) was sampled at two different stages: During transplantation (30 DAS) and at maturity stage (135 DAS). Soil samples were obtained using PVC cores/pipes of 25 cm in length at each of these sampling points. The samples were subsequently air-dried, ground, passed through a 5 mm nylon sieve, and homogenized to create representative samples. The soil's physicochemical and biological properties, such as pH, electrical conductivity (EC), redox potential (ORP), and soil organic carbon (SOC), were analyzed using appropriate standard methods as detailed in Upadhyay et al. ( 2021 ). Harvesting, Growth Measurement, and Yield Evaluation Plant sampling was done at two key time points throughout the cultivation period. The first sampling was done at 65 DAT, during the milking stage of cultivation. At this stage, the leaf samples were collected for the estimation of chlorophyll content and other biochemical assays. The second sampling was conducted at maturity (135 DAS). During this stage, root and shoot lengths were measured, spikes were separated, and fertile spikelet per spike were counted. Grains were separated, dried, and the weight of 1000 grains was determined to estimate the yield. Measurement of Electrolytic Leakage and Lipid Peroxidation Electrolytic leakage (EL) was evaluated according to the protocol of Dionnisio-Sese and Tobita (1998). Fresh leaf samples (20 leaf discs) were submerged in 40 ml of deionized water, and after 10 minutes, electrical conductivity (EC1) was recorded. The solution was then subjected to controlled temperature baths at 50–60°C for 25 minutes, and EC2 was measured. Lastly, the solution was boiled at 100°C for 10 minutes, and EC3 was recorded. Electrolytic leakage was calculated using the formula: Electrolytic leakage (%) = \(\:\frac{(EC2-EC1)}{EC3}\times\:100\) Lipid peroxidation was measured by estimating the production of malondialdehyde (MDA) using the protocol of Heath and Packer ( 1968 ). Measurement of Pigments Pigments, including total chlorophyll and carotenoids, were assessed from fresh leaf samples using chilled acetone extraction and spectrophotometric measurements. Chlorophyll and carotenoid concentrations were determined through established methodologies as outlined by Arnon ( 1949 ) for chlorophyll and Duxbury and Yentsch ( 1956 ) for carotenoids. Leaf samples weighing between 20–25 mg were finely crushed and homogenized with 1 ml of 80% acetone, followed by centrifugation at 10,000 rpm for 15 minutes. The resulting supernatant was then employed for the analysis of chlorophyll a, chlorophyll b, and carotenoids, with absorbance readings taken at 480, 510, 645, and 663 nm using a UV-VIS spectrophotometer (Model-UV-1900, Shimadzu, India). Subsequent to the absorption measurements, the values were computed using specific calculations. Protein content determination and antioxidant enzyme assays Protein content determination utilized the Bradford ( 1976 ) method. In this approach, a 1 ml sample, comprising 950 µl Bradford reagent and 50 µl enzyme extract, was meticulously mixed using a vortex. Following 10-minute incubation, the solution underwent a discernible transformation into a blue color, and the absorbance was measured at 650 nm. This method capitalizes on the distinctive color change at the specified wavelength for precise assessment of protein content. The activities of antioxidant enzymes superoxide dismutase (SOD), guaiacol peroxidase (GPX), ascorbate peroxidase (APX) and catalase (CAT) were assayed by following the methods of Beauchamp and Fridovich ( 1971 ), Hemeda and Klein ( 1990 ), Nakano and Asada (1981) and Aebi (1974) as detailed previously (Srivastava et al., 2006). Arsenic analysis The plant samples were separated into different parts (root, flag leaves, grains). These parts were cleaned by gentle shaking and placed into beakers for oven-drying. Dried samples were collected, and grain samples were dehusked to get the edible grain samples and crushed into powder form. The samples were digested with HNO3 (70% Suprapur Merck; Darmstadt-Germany) in a heating test tube block digester at 180°C for 1 hour and then at 200 o C for 45–60 minutes till the sample turned into a white crystalline appearance (Srivastava et al., 2014 ; Pathak et al., 2024 ). After proper digestion, the final volume for each sample was maintained up to 10 ml with ultrapure (Type I) water (Milli-Q) and filtered with a 0.22 µm pore size filter paper. Finally, the P and As contents were measured through (HG-AAS). For hydride generation, analytical standard sodium borohydride (3%; Merck), sodium hydroxide (2.5%; Merck) and hydrochloric acid (6 M; Merck) were used. The precision of the analysis was checked by certified standard reference materials (SRMs) (NIST, USA), such as 1568a rice flour and 1573a tomato leaf, as per the details given in Khanam et al. ( 2022 ). The quality control was performed by triplicate analysis and by the measurement of the recovery of spiked digested samples. The recoveries varied from 95 to 98% for all the tested samples. Statistical Analysis The experiment was conducted using a Randomized Complete Block Design (RCBD) with three replications. Data were subjected to two-way analysis of variance (ANOVA) to evaluate the effects of location and treatment, as well as their interaction. When significant differences were detected, mean comparisons were performed using Tukey’s Honestly Significant Difference (HSD) test at a probability level of p ≤ 0.05. Statistical analyses were carried out using GraphPad Prism. Results Soil physicochemical characteristics and soil enzymes The soil samples were collected at two different times, and variations were observed in the blocks of both varieties under different P doses. The analysis of soil samples from both sites, represented in Supplementary Tables S1 and S2, revealed significant changes in various soil parameters. A decrease in pH was noticed from sampling at the vegetative stage to the heading phase, while EC was increased at both sites. There were some changes in ORP, soil organic carbon and enzymatic activities also. Soil As levels were also examined, showing changes from the initial concentrations for all treatments at both sites (supplementary Table S1 and S2). The level of soil As was generally decreased from vegetative to heading phase, and this decline was greater for SK blocks than for the DS blocks. In contrast to As, soil P levels were found to increase from the vegetative to the heading phase. These findings underscore the dynamic nature of soil properties and enzymatic activities in response to varying P doses, highlighting the importance of monitoring and understanding soil dynamics for sustainable agricultural practices. Morphological differences in selected varieties The influence of different levels of phosphorus fertilizer on the physical characteristics and yield of two varieties was noticeable. Both varieties showed a decline in their growth and yield when given only half the recommended P dose, while they displayed an increase with a high P dose. The SK variety experienced a more significant decrease. The DS variety showed better changes in root structure, with a 19% and 18% increase in length at Ghazipur and Jaunpur, respectively, with high P doses (Fig. 1 a). In contrast, the SK variety showed a 14% and 3% increase at the same sites at high P dose. SK also experienced a greater decrease in shoot length, with reductions of 14% and 35% under low P doses at the respective sites, while DS showed a smaller decrease (Fig. 1 b). Panicle length decreased notably for the SK variety under low P doses, while DS showed minimal reductions. The number of fertile spikelet decreased under low P doses, impacting grain weight and number per panicle. SK, being less efficient in P utilization, showed a more significant overall reduction. However, both varieties responded positively to increased P doses, leading to improved production and yield at both sites (Fig. 1 a-f). Effect on electrolytic leakage and MDA Malondialdehyde, an indicator of membrane damage, was used to assess the extent of stress under low P doses. MDA production in roots increased by 23% and 18% for SK, and 7% and 8% for DS at sites 1 and 2, respectively, under low P doses as compared to the control (Fig. 2 a). In shoots, MDA increased by 17% and 11% for SK, and 6% and 2% for DS at the respective sites. However, at high P doses at both sites, either a slight increase or no significant change in MDA was noticed in both SK and DS. At P2 in Ghazipur, shoots of both SK and DS showed a significant decline in MDA than the control (Fig. 2 a-b). Hence, high P dose proved to have an ameliorative effect. Plants subjected to a reduced P fertilizer dose also exhibited elevated electrolytic leakage (EL) in both roots and shoots, indicating heightened stress levels. Root EL increased by 27% and 8% for SK, and 10% and 2% for DS at sites 1 and 2, respectively, compared to the control dose. Similarly, shoot EL increased by 23% and 19% for SK, and 5% and 14% for DS at sites 1 and 2, respectively (Fig. 2 c-d). The rise in EL suggests compromised membrane integrity in response to nutrient stress. However, when plants received high P fertilizer supply, double the recommended dose, stress levels were reduced in both varieties compared to the half P and normal P doses (Fig. 2 c-d). Effect on photosynthetic pigments Phosphorus, as a vital nutrient, plays a crucial role in the process of photosynthesis within plants. The deficiency of this essential nutrient can significantly impede the photosynthetic process. In our experiment, the supplementation of a low P dose resulted in a notable decrease in pigment levels, with a more pronounced effect being in the SK variety. The reduction in pigment content was statistically significant, revealing a decrease in chlorophyll a (chl a) by 29% and 27% for SK, and 14% and 18% for DS at sites 1 and 2, respectively. Chlorophyll b (chl b) exhibited reductions of 18% and 13% for SK, and 8% and 10% for DS at the respective sites, while carotenoid content decreased by 35% and 32% for SK, and 21% and 14% for DS at sites 1 and 2, respectively, compared to the control (Fig. 3 a-c). Conversely, under conditions of high P dose, an increase in chl a, chl b, and carotenoid content was observed in comparison to the control. The increment in chl a was evident in both varieties, with a rise of 10% and 15% for SK, and 13% and 18% for DS at sites 1 and 2, respectively. Chl b showed increases of 30% and 27% for SK, and 34% and 33% for DS at the respective sites (Fig. 3 a-c). These findings underscore the pivotal role of phosphorus in sustaining optimal pigment production, ultimately influencing the efficiency of the photosynthetic process in plants. Responses of antioxidant enzymes (SOD, APX, CAT and GPX) Superoxide dismutase (SOD) activity was assessed in the roots and shoots of both selected plant varieties, revealing noteworthy variations in response to phosphorus (P) fertilizer levels. SOD activity exhibited a reduction under a low P dose. The reduction in SOD activity under low P doses was 11% and 24% for SK, and 4% and 10% for DS at sites 1 and 2, respectively. Similarly, in the shoots, a reduction in SOD activity was noted, which was 22% and 31% for SK, and 13% and 18% for DS at sites 1 and 2, respectively, compared to the control. Conversely, when plants received an adequate supply of P fertilizer, an increase in SOD activity was observed. In the roots, SOD activity rose by 9% and 36% for SK, and 9% and 25% for DS at sites 1 and 2, respectively. In the shoots, the increase in SOD activity was recorded as 17% and 16% for SK, and 5% and 8% for DS at sites 1 and 2, respectively, compared to the control dose. These findings highlight the responsive nature of SOD activity to varying P fertilizer levels (Fig. 4 a-b). A consistent trend was observed for APX activity, mirroring the patterns seen with SOD, in response to varying P fertilizer levels. APX activity in the roots and shoots exhibited a decrease when compared to the control dose for both varieties, with reductions being greater for SK than for DS at both sites (Fig. 4 c-d). In contrast, under higher P doses, APX activity increased for both varieties at both sites. This rise in APX activity suggested a positive response to an increased supply of phosphorus, indicating the potential role of phosphorus in enhancing the antioxidant defense mechanisms mediated by APX. The activity of the CAT enzyme showed a reduction in the roots under a half P dose, with decreases of 25% and 8% for SK, and 20% and 12% for DS at sites 1 and 2, respectively. In shoots, CAT activity declined by 20% and 29% for SK, and 5% and 13% for DS at sites 1 and 2, respectively (Fig. 5 a-b). Under a double P dose, CAT activity increased by 8% at site 1 and 6% at site 2 for DS, while for SK, there was an exceptional decrease of 6% at site 1 and an increase of 15% at site 2 (Fig. 5 a-b). The decline in CAT activity was notable under a half P dose compared to the control, further emphasizing the impact of P availability on antioxidant enzyme activity in plants. GPX activity in the roots exhibited a decline under a half phosphorus dose, with reductions of 30% and 13% for SK, and 1% and 3% for DS at sites 1 and 2, respectively. Similarly, in the shoots, GPX activity decreased by 13% for SK at site 1 and a significant 35% at site 2. For DS, there was a 10% increase at site 1, but a reduction of 6% at site 2 (Fig. 5 c-d). Conversely, under double P dose, both root and shoot GPX expression increased for both varieties at both experimental sites, suggesting a positive response to higher P levels. Phosphorus and arsenic accumulation pattern in rice The study revealed notable fluctuations in P accumulation patterns in response to varying doses of P fertilizer at two distinct sites, Ghazipur and Jaunpur. Alterations in P concentrations were observed across different plant parts, including the root, shoot, and grains, highlighting the responsiveness of these components to changes in P fertilizer dosage. The SK variety exhibited a significant reduction in P concentration; for example, a 17% decrease in root P accumulation under half-dose conditions at the Ghazipur site. Conversely, the DS variety displayed a non-significant reduction of only 3% in root P accumulation under similar conditions. Upon application of a double dose of P fertilizer, root P accumulation increased by 15% for the DS variety and 8% for the SK variety at site 1, with corresponding increases of 3% and 10% being noticed at site 2 (Fig. 6 a-c). Significant differences were observed in shoot and grain P accumulation for both varieties under varying P fertilizer doses. Under half-dose conditions, the SK variety experienced a substantial decrease of 21% and 20% in shoot P accumulation at the Ghazipur and Jaunpur sites, respectively, while the DS variety exhibited a negligible reduction. In terms of grain P accumulation, the DS variety demonstrated higher accumulation in both control and double P dose conditions, with negligible reduction under half-dose conditions. In contrast, the SK variety exhibited a noticeable reduction of 18–20% in P accumulation for both sites under similar conditions (Fig. 6 a-c), highlighting varietal disparities in the response to P fertilizer doses across different plant parts. Arsenic concentration within both the vegetative parts and grains exhibited sensitivity to variations in P fertilizer dosage. Under a low P dose, there was an elevation in As accumulation, whereas the addition of a double P dose resulted in a slight decrease. Notably, the accumulation of As was consistently higher in SK tissues and grains across all conditions, with an enhanced accumulation observed particularly under low P doses. In SK grains, the As concentration in control conditions was recorded at (0.58–0.6 mg kg − 1 ). Under half the standard P dose, a significant increase to (0.69–0.75 mg kg − 1 ) was observed, indicating about a 20% rise at both contaminated sites. Conversely, applying a double P dose led to a reduction in As accumulation by 9–11% at both sites (Fig. 7 a-c). In the case of the DS variety, the tissue and grain arsenic accumulation demonstrated resilience to lower P doses, with minimal effects. However, an application of higher P doses resulted in a significant reduction of As accumulation (Fig. 7 a-c). These findings underscore the differential response of arsenic accumulation in vegetative tissues and grains of SK and DS varieties to varying P fertilizer levels, offering valuable insights into the intricate interactions between phosphorus and arsenic dynamics in agricultural systems. Rice yield of two cultivars grown in different fields under different P fertilizer doses The assessment of rice yield, conducted within 100 ft 2 plots, revealed a range spanning from 5683 to 6496 kg ha − 1 , as delineated in Table 1 . A significant reduction in yield was seen under conditions of low P dosage. Specifically, at the Jaunpur site, the SK variety exhibited a substantial 31% decline, with yield plummeting from 3637 to 2492 kg ha − 1 . Conversely, the DS variety demonstrated a lesser reduction of 15% under analogous conditions. Parallel observations were made at the Ghazipur site, where the SK and DS varieties experienced reductions of 29% and 12%, respectively under low P (Table 1 ). Under high P, an increase in yield was observed for both varieties in both sites; the increase in yield ranged from 12–14% for SK and 17–20% for DS (Table 1 ). Table 1 Yield parameters of rice varieties. SK Variety DS Variety Yield (kg/100ft 2 ) Yield (kg/h) Yield (kg/100ft 2 ) Yield (kg/h) Jaunpur Control 3.38 ± 0.31 3637 ± 329.71 3.88 ± 0.22 4180 ± 235.78 P1 2.32 ± 0.21 2492 ± 230.37 3.29 ± 0.25 3537 ± 268.72 P2 3.86 ± 0.23 4157 ± 250.30 4.65 ± 0.24 5007 ± 255.16 Ghazipur Control 2.95 ± 0.13 3179 ± 141.06 3.38 ± 0.42 3641 ± 456.77 P1 2.08 ± 0.16 2241 ± 177.11 2.99 ± 0.41 3221 ± 445.21 P2 3.31 ± 0.29 3568 ± 309.77 3.94 ± 0.46 4238 ± 496.93 Discussion Rice ( Oryza sativa L.), an essential food item for over half of the global population, is grown in over 100 countries, of which close to 90% of its total production is focused on the continent of Asia. Owing to its high calorific value, economic affordability, bland flavour, high iron content, and relatively low potential for allergenicity, rice forms an ideal constituent of the diet, particularly of children (Hoang et al., 2022). Arsenic (As) toxicity offers a two-sided threat by posing a threat not only towards the production of agriculture globally but also by being an effective human carcinogen. It is therefore essential that the problem be dealt with by improving crop sustainability as well as public health (Banerjee et al., 2023 ). The results of the present study propose that the DS variety exhibited superior performance in response to varying P fertilizer doses compared to the SK variety. Several key factors highlight the resilience and adaptability of the DS variety. Firstly, in terms of morphological attributes, the DS variety demonstrated better root and shoot growth under low P conditions, with a substantial increase in root length compared to SK (Fig. 1 a-b). This superior root development may confer greater tolerance to nutrient stress, enabling DS plants to access and absorb nutrients more efficiently, ultimately contributing to enhanced overall growth even when the P is in scarcity. The evolution of higher plants has led to the development of various strategies for acquiring nutrients from the soil, especially for less available elements like P (Shen et al., 2011 ). One notable adaptation involves long root hairs at high densities in the roots of Arabidopsis plants. This adaptation allows plants to effectively explore a larger volume of soil and capture more nutrients (Lynch 2019 ). Thus, amended root growth is connected to better P nutrition of plants. Additionally, the DS variety unveiled more favourable responses in terms of shoot length, panicle length, and fertile spikelet development under varying P doses. The increased shoot and panicle lengths (Fig. 1 c-f) suggest a greater above-ground growth, which is crucial for maximizing photosynthetic potential and overall biomass production. Moreover, the higher number of fertile spikelets indicates better reproductive success, contributing to potential increases in yield (Table 1 ) (Pretini et al., 2021 ; Diarra et al., 1985 ). Plant breeders often focus on optimized panicle structures for developing rice varieties with enhanced yield potential (Sabar et al., 2024 ). Previous studies on soybean, the effect on growth and yield was highly significant (α < 0.05) with an increase in root/shoot dry and fresh weight in plants when a mixture of Bradyrhizobium inoculums along with phosphorus was used (Rabbani et al., 2023 ; Mirriam et al., 2023 ). Soil application of phosphorus resulted in a noteworthy rise in both yield and growth index in maize (Khaleeq et al., 2023). When initial phosphorus deficiency symptoms manifested 25 days after sowing in wheat, the application of higher doses of ammonium phosphate via foliar spray demonstrated the most significant reduction in phosphorus deficiency and led to the highest yields (Haloi, 1980 ). Thus, previous reports also support the contemporary idea. Figure 8 summarizes the mechanism by which excess P lessens As toxicity in rice grown in As contaminated soil. In the present study the morphological changes were conveyed by alterations in physiological processes. The grade of cell membrane damage (lipid peroxidation of membrane) and ROS buildup was determined by MDA and H 2 O 2 quantification, and their pattern was consistent with cell death. Excessive production of ROS upon arsenic stress led to oxidation of cellular as well as organelle membrane lipids and MDA production. ROS triggers membrane lipid peroxidation by withdrawing an electron from the hydrogen atom of the fatty acyl chain of PUFA (polyunsaturated fatty acid), which makes the membrane rigid, leaky, and damaged membrane proteins (Abbas et al., 2018 ). Disrupted membranes result in leakage of crucial components such as electrolytes from the cells, eventually leading to cell death (Salam et al., 2022 ). In the present work, MDA accumulation reflected considerable differences between varying plant types as well as different levels of P fertilization. At low P availability, roots and shoots of SK reflected a noticeable elevation of MDA concentrations compared with DS, which shows that SK is rather more susceptible to oxidative damage. The higher MDA concentrations linked with P deficiency confirm increased production of reactive oxygen species (ROS) and lower membrane stability. By contrast, DS reflected relatively lower MDA concentrations that suggest higher inherent tolerance as well as more effective management of oxidative stress when faced with P limitations (Fig. 2 a-d). Such genotypic differences in oxidative damage have been thoroughly reported for cereal species treated with nutrient stress regimes. Notably, at high P treatment doses, MDA concentrations either stayed the same or decreased, specifically at P2 in Ghazipur for shoots, wherein SK and DS significantly lowered from controls. This outcome indicates the beneficial action of sufficient P availability toward sustaining cellular redox balance as well as towards lowering membrane lipid peroxidation. Sufficient phosphorus is well documented for increased energy metabolism, enhanced nucleic acid and protein synthesis, and enhanced photosynthetic potential that mitigates ROS accumulation. This idea also finds corroboration from Al-Qahtani et al. (2024). Overall, these results underscore a bivalent role of phosphorus hereafter: deficiency of phosphorus causes oxidative stress as indicated by increased MDA accumulation, while sufficient or higher P supply mitigates oxidative membrane damage by enhancing enzymatic antioxidant defense. As per reports, photosynthetic activity is inhibited by arsenic tremendously due to the shifting that occurs in light-harvesting complexes that involve pigment losses as well as chlorophyll production. The Mg-ion of chlorophyll is replaced by As, disrupting photosynthesis (Banerjee et al., 2024 ). Figure 3 clearly depicts that reduced P doses hampered pigment production, leading to a substantial decrease in chlorophyll a, chlorophyll b, and carotenoid levels, with the SK variety experiencing more pronounced reductions than DS due to As toxicity. However, an adequate P supply mitigated these effects, resulting in increased pigment production by upregulating essential nutrient uptake. It is universally accepted that ROS hampers the overall health and productivity of plants. Superoxide dismutase (SOD), alongside glutathione peroxidase (GPX), is one of the key enzymatic antioxidants that scavenge ROS and cause subsequent alleviation of oxidative stress (Ighodaro and Akinloye, 2017). The results (Figs. 4 and 5 ) clearly demonstrate that the antioxidant enzymatic activities of rice are strongly regulated by the availability of phosphorus (P). Consistent decline of SOD, APX, CAT, and GPX activities by half P rates indicates that the plant ROS-scavenging system is influenced by nutrient limitation, thereby enhancing susceptibility to oxidative stress. However, adequate P supply overcame and, for most instances, enhanced such enzymatic activities, indicative of adequate P fertilization sustaining redox homeostasis as well as stress resistance (Khan et al., 2023 ). Among the enzymes studied, SOD and APX possessed the highest consistent positive response against higher P levels, which indicates their central role in the first defense against oxidative stress (Fig. 4 a-c). CAT and GPX were more variable, particularly by site and variety, which reflects variation in their regulatory mechanisms. SK also responded more to P variation than did DS, which indicates genotypic variation of antioxidant controls (Fig. 5 a-d). The results are in line with Mishra et al. ( 2022 ), which showed similar evidence. In summary, varietal variation indicates potential for P-efficient genotypes possessing enhanced resistance against nutrient stress and provides useful insights towards sustainable nutrient use in rice production. Some other studies showed that elevated P level has an ameliorative effect against various HM stress and influence plant growth and development. Arshad et al., ( 2016 ) reported the provision of P enhanced shoot biomass, leaf area, photosynthetic pigments, and mineral nutrient levels, while decreasing concentrations of cadmium (Cd) and hydrogen peroxide (H 2 O 2 ) in wheat shoots. The application of P also improved antioxidant enzyme activities and gas exchange characteristics, serving as a crucial mechanism for enhancing Cd tolerance in wheat. A study conducted by Tairo et al. (2013) demonstrated that the addition of P led to a significant increase in growth attributes of soybean, such as the leaf area index, improved efficiency in photosynthesis, and promoted green pigment formation. In addition to biochemical and morphological changes, the major finding of the present study was the decline in As content of plant tissues, including grains, along with an enhancement of P content in high P supply. The research disclosed considerable varietal variation in P accumulation of different plant organs in response to different fertilizations. The SK variety was very sensitive to phosphate starvation, as evidenced by considerable decreases in P concentrations of root, shoot, and grains when they were supplied from half-doses during both locations of experiments. In contrast, the DS variety registered marginal decreases in root and shoot P contents while sustaining grain P accumulation, suggesting high tolerance to restricted phosphate supplies. Under double-dose fertilization, both genotypes registered greater P accrual, but DS recorded relatively larger increments in root and grain P compared with SK, suggesting greater utilization and acquisition efficiency. DS tended to maintain consistent P allocation across the root, shoot, and grain, while there was strong dependence on high external P application in SK. The study results (Fig. 6 ) showed that DS has improved adaptability to variable phosphorus availabilities, hence making it more suitable for farming on phosphorus-deficient soils. On the contrary, SK might require fertilizer management schedules tailor-made to sustain yield levels. The reduction in As content in roots, shoots, and grains could be attributed to the competition between As and P for the same transporters and binding sites. Arsenic enters rapidly across the root cells through phosphate transporters (PHTs) are characterized through their sequence homology with inorganic phosphate (Shukla et al., 2024 ; Banerjee et al., 2024 ). Hence, optimizing P nutrition to rice plants appears to be a viable and practical approach to mitigate As toxicity and reduce its accumulation in grains. Importantly, the reduction in As accumulation also resulted in higher yields of rice in high P supply. Hence, optimal P nutrition is a requisite of the plant that, when fulfilled, makes the plant grow and produce better. The present study also highlighted the importance of genetic variability in plant responses to nutrient stress (Basavaraj et al., 2022 ). The study emphasizes the importance of selecting crop varieties with inherent traits that contribute to better nutrient utilization and stress tolerance. In conclusion, the findings of this study, coupled with relevant references, provide substantial evidence supporting the superiority of the DS variety under varying phosphorus conditions. This underscores the significance of genetic diversity in crop breeding programs and the potential for selecting varieties that can thrive under suboptimal nutrient conditions, contributing to sustainable and resilient agriculture. Conclusion This study clearly establishes that phosphorus nutrition plays an overriding role in regulating arsenic dynamics, stress physiology, and yield performance in arsenic-contaminated soil-grown rice. Out of two cultivars tested, which one exhibited high and low phosphorus use efficiency (PUE), respectively, the high PUE variety DNA Sribala (DS) showed, on average, higher tolerance to nutrients as well as arsenic toxicity than the low PUE variety Sai Kasturi (SK). The increased tolerance exhibited by DS was expressed through greater root and shoot growth, higher panicle length, and higher number of fertile spikelets, and these, in totality, resulted in more stable and reproducible yield outcomes, even under low phosphorus conditions. Physiological and biochemical studies also showed that DS retained higher contents of photosynthetic pigments and possessed a better antioxidant defense mechanism, reflected in increased SOD, APX, GPX, and CAT activities under increased P supply. These adjustments effectively controlled oxidative damage, as reflected in decreased production of MDA and electrolyte leakage. Notably, P supplementation not only alleviated stress-caused damage but also substantially diminished As deposition in vegetative organs and grains, especially at elevated P doses. Though SK showed increased susceptibility to both low P and arsenic toxicity, DS showed an even and stable nutrient uptake profile, supporting the importance of P-efficient genotypes in contaminated agroecosystems. In short, this study shows double benefits: reducing arsenic seepage into the food supply and maximizing rice production. These results underscore the importance of precision nutrient management and judicious varietal selection as an environmentally sustainable agricultural practice to safeguard vulnerable areas threatened by arsenic. However, further studies are necessary to confirm these results under various soils and agro-climatic regimes to understand P-As interaction. Molecular research determine genes responsible for phosphorus use efficiency and arsenic tolerance would support breeding programs to develop resilient varieties. Integrating P-efficient varieties with superior nutrient management practices involving biofertilizers or nano fertilizers would also portray increased productivity and lower arsenic uptake. These measures, in conjunction with location-specific management guidelines, will be essential to provide crop safety and sustainable rice production in arsenic-contaminated areas. Declarations Acknowledgements The authors are thankful to IESD-BHU, Varanasi, India, for the lab facility. SS is thankful to IOE-BHU (No.-6031) for financial support. KS is thankful to CSIR for Senior Research Fellowship (09/013(0818)/2018-EMR-I). Author contributions KS executed all the experiments, formal analysis, and data curation, and prepared the draft the SKP helped in formal analysis and writing the original draft. SB helped in writing the original draft. SS conceptualized and supervised the study and finalized the draft for submission. Availability of data and materials The data that support the findings of this study are available from the corresponding author upon reasonable request. Declaration of interests The authors declare no conflict of interest. References Abbas, G., Murtaza, B., Bibi, I., Shahid, M., Niazi, N. K., Khan, M. I., … Natasha. (2018). Arsenic uptake, toxicity, detoxification, and speciation in plants: physiological, biochemical, and molecular aspects. International journal of environmental research and public health , 15 (1), 59. Aebi, H. (1974). Catalase. In Methods of enzymatic analysis (pp. 673–684). Academic press. Nakano, Y., & Asada, K. (1981). 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Supplementary Files KavitaSaurabhFinalMSHEPSupplementaryfileV2.docx KavitaSaurabhFinalMSHighlightsV2.docx GA.png Graphical Abstract Cite Share Download PDF Status: Under Review Version 1 posted Editorial decision: Revision requested 27 Feb, 2026 Reviewers agreed at journal 25 Feb, 2026 Reviews received at journal 24 Feb, 2026 Reviews received at journal 24 Feb, 2026 Reviews received at journal 22 Feb, 2026 Reviewers agreed at journal 21 Feb, 2026 Reviewers agreed at journal 21 Feb, 2026 Reviewers agreed at journal 21 Feb, 2026 Reviewers agreed at journal 19 Feb, 2026 Reviewers agreed at journal 19 Feb, 2026 Reviewers agreed at journal 19 Feb, 2026 Reviewers agreed at journal 19 Feb, 2026 Reviewers invited by journal 19 Feb, 2026 Editor assigned by journal 18 Feb, 2026 Submission checks completed at journal 18 Feb, 2026 First submitted to journal 16 Feb, 2026 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. 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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-8891770","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":595047947,"identity":"fa976378-e84e-457c-8b05-2d56816da1d3","order_by":0,"name":"Kavita Shukla","email":"","orcid":"","institution":"Banaras Hindu University","correspondingAuthor":false,"prefix":"","firstName":"Kavita","middleName":"","lastName":"Shukla","suffix":""},{"id":595047948,"identity":"d2e6a53c-4ff3-4167-a342-42d4eeba07bd","order_by":1,"name":"Saurabh Kumar Pathak","email":"","orcid":"","institution":"Banaras Hindu University","correspondingAuthor":false,"prefix":"","firstName":"Saurabh","middleName":"Kumar","lastName":"Pathak","suffix":""},{"id":595047949,"identity":"c697c789-6391-4e77-98ef-63db86e699f3","order_by":2,"name":"Swarnendra Banjerjee","email":"","orcid":"","institution":"Southern Federal University","correspondingAuthor":false,"prefix":"","firstName":"Swarnendra","middleName":"","lastName":"Banjerjee","suffix":""},{"id":595047950,"identity":"c3051436-7e91-45aa-bcb7-e9f40bcb5b44","order_by":3,"name":"Sudhakar Srivastava","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA9UlEQVRIiWNgGAWjYHACMwaGA0CKGYh5Kg4kQAQNiNZyhiQtIMDbBtOCB8jPSN724McZu3wGdubDH97Ou5NncID54QeGgjs4tRjcSCs37LmRbNnAzJYmOXfbs2KDA2zGEgwGz3Brkcgxk+D5wGzAwMxjxsy77XDihgMgpxocxuOwHDPJPx/qgVr4P3/mnQPSwv4NrxaGGzlm0jw3DoNsYZDmbQBp4cFvi8GZZ2XSMmeOG7Axs5lJzjn2LHHmYZ5iiQR8DmtP3ib55li1AT//4ccf3tTcSew73r7xw4c/eBwGA2xwFihOEwhrGAWjYBSMglGABwAA1C5VVe9ke3sAAAAASUVORK5CYII=","orcid":"","institution":"Banaras Hindu University","correspondingAuthor":true,"prefix":"","firstName":"Sudhakar","middleName":"","lastName":"Srivastava","suffix":""}],"badges":[],"createdAt":"2026-02-16 09:53:34","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-8891770/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-8891770/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":103253250,"identity":"c9971681-935a-4c7e-b77d-e8adb325b3d4","added_by":"auto","created_at":"2026-02-23 16:21:11","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":65393,"visible":true,"origin":"","legend":"\u003cp\u003eFigure (a-f) showing the morphological appearance, root length (cm), shoot length (cm), panicle length and number of fertile spiklets, thousands grain weight (g), number of grains per spikelet under control (C), half (P1) and double (P2) phosphorus fertilizer dose for selected variety (SK, DS) at the respective As contaminated sites (Ghazipur and Jaunpur). Bars represent mean values. Error bars indicate ± standard error (SE). Different lowercase letters above bars indicate significant differences among treatments within each location at P ≤ 0.05 (Two-way ANOVA followed by Tukey’s HSD test).\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-8891770/v1/9268f9b5795f78952179f8a2.png"},{"id":103253251,"identity":"a002aa82-8e26-415a-8c7e-2879dd40457c","added_by":"auto","created_at":"2026-02-23 16:21:11","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":79440,"visible":true,"origin":"","legend":"\u003cp\u003eFigure (a-d) showing the physiological response (a-root MDA, b- shoot MDA, c- root electrolytic leakage, d- shoot electrolytic leakage) under control (C), half (P1) and double (P2) phosphors fertilizer dose for selected variety (SK, DS) at the respective As contaminated sites (Ghazipur and Jaunpur). Bars represent mean values. Error bars indicate ± standard error (SE). Different lowercase letters above bars indicate significant differences among treatments within each location at P ≤ 0.05 (Two-way ANOVA followed by Tukey’s HSD test).\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-8891770/v1/d2ac28b37bbd433d366acb33.png"},{"id":103506078,"identity":"3d486ba1-97ff-4cb5-99b3-c0e6e5f55481","added_by":"auto","created_at":"2026-02-26 13:34:00","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":63787,"visible":true,"origin":"","legend":"\u003cp\u003eFigure (a-c) showing the physiological response (a-chl a, b-chl b, c- carotenoids) under control (C), half (P1) and double (P2) phosphorus fertilizer dose for selected variety (SK, DS) at the respective As contaminated sites (Ghazipur and Jaunpur). Bars represent mean values. Error bars indicate ± standard error (SE). Different lowercase letters above bars indicate significant differences among treatments within each location at P ≤ 0.05 (Two-way ANOVA followed by Tukey’s HSD test).\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-8891770/v1/96e738b6e50efe68eb3d3984.png"},{"id":103505121,"identity":"ac0f8a4d-876e-4055-80da-a38f1c3a1d93","added_by":"auto","created_at":"2026-02-26 13:24:30","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":75649,"visible":true,"origin":"","legend":"\u003cp\u003eFigure (a-d) showing the biochemical response (a-root SOD, b-shoot SOD, c-root APX, d-shoot APX) under control (C), half (P1) and double (P2) phosphorus fertilizer dose for selected variety (SK, DS) at the respective As contaminated sites (Ghazipur and Jaunpur). Bars represent mean values. Error bars indicate ± standard error (SE). Different lowercase letters above bars indicate significant differences among treatments within each location at P ≤ 0.05 (Two-way ANOVA followed by Tukey’s HSD test).\u003c/p\u003e","description":"","filename":"4.png","url":"https://assets-eu.researchsquare.com/files/rs-8891770/v1/1431cf3d2fbe222a8a5a5f13.png"},{"id":103253260,"identity":"b7f7ae22-b636-4f03-8648-b0d393dc0c43","added_by":"auto","created_at":"2026-02-23 16:21:12","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":74679,"visible":true,"origin":"","legend":"\u003cp\u003eFigure (a-d) showing the biochemical response (a-root GPX, b-shoot GPX, c-root CAT, d-shoot CAT) under control (C), half (P1) and double (P2) phosphorus fertilizer dose for selected variety (SK, DS) at the respective As contaminated sites (Ghazipur and Jaunpur). ). Bars represent mean values. Error bars indicate ± standard error (SE). Different lowercase letters above bars indicate significant differences among treatments within each location at P ≤ 0.05 (Two-way ANOVA followed by Tukey’s HSD test).\u003c/p\u003e","description":"","filename":"5.png","url":"https://assets-eu.researchsquare.com/files/rs-8891770/v1/ddacfdceb10ece5121abfa4f.png"},{"id":103253252,"identity":"f0339c37-a84d-44db-aeaa-7084fa49810f","added_by":"auto","created_at":"2026-02-23 16:21:11","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":53145,"visible":true,"origin":"","legend":"\u003cp\u003eFigure (a-c) showing the P accumulation (a root P concentration, b- shoot P concentration, c-grain As concentration) under control, half and double dose of P fertilizer for selected variety (DS and SK). Bars represent mean values. Error bars indicate ± standard error (SE). Different lowercase letters above bars indicate significant differences among treatments within each location at P ≤ 0.05 (Two-way ANOVA followed by Tukey’s HSD test).\u003c/p\u003e","description":"","filename":"6.png","url":"https://assets-eu.researchsquare.com/files/rs-8891770/v1/dbeb827ec0a0c4ac78956901.png"},{"id":103253254,"identity":"7f08044d-db8f-4669-bd61-d39acb66224c","added_by":"auto","created_at":"2026-02-23 16:21:11","extension":"png","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":48807,"visible":true,"origin":"","legend":"\u003cp\u003eFigure (a-c) showing the As accumulation (a root As concentration, b- shoot As concentration, c-grain As concentration) under control, half and double dose of P fertilizer for selected variety (DS and SK). All the values are means of triplicate ±SD. Bars represent mean values. Error bars indicate ± standard error (SE). Different lowercase letters above bars indicate significant differences among treatments within each location at P ≤ 0.05 (Two-way ANOVA followed by Tukey’s HSD test).\u003c/p\u003e","description":"","filename":"7.png","url":"https://assets-eu.researchsquare.com/files/rs-8891770/v1/6cf2ec394fb98ea57e579249.png"},{"id":103253259,"identity":"476b68ee-a236-4b08-9baa-f7ce460c1eb4","added_by":"auto","created_at":"2026-02-23 16:21:11","extension":"png","order_by":8,"title":"Figure 8","display":"","copyAsset":false,"role":"figure","size":313594,"visible":true,"origin":"","legend":"\u003cp\u003eSchematic representation shows arsenic toxicity rice and its mitigation through using excess phosphorous in arsenic-contaminated soil.\u003c/p\u003e","description":"","filename":"8.png","url":"https://assets-eu.researchsquare.com/files/rs-8891770/v1/954cef32883db46f24f8b456.png"},{"id":104397642,"identity":"5af00a69-d04a-427e-bb6a-23fa595cfb24","added_by":"auto","created_at":"2026-03-11 11:53:41","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1497267,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8891770/v1/338c8cb3-ac4f-4e65-a7d4-3f181a04ff58.pdf"},{"id":103505164,"identity":"274012b6-3626-4f7b-826f-5849891cdf86","added_by":"auto","created_at":"2026-02-26 13:25:51","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":236316,"visible":true,"origin":"","legend":"","description":"","filename":"KavitaSaurabhFinalMSHEPSupplementaryfileV2.docx","url":"https://assets-eu.researchsquare.com/files/rs-8891770/v1/35598fea99e7b1ac7d2885a3.docx"},{"id":103253257,"identity":"12a2dd4d-65bf-4a34-a2e6-95bec6437624","added_by":"auto","created_at":"2026-02-23 16:21:11","extension":"docx","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":16985,"visible":true,"origin":"","legend":"","description":"","filename":"KavitaSaurabhFinalMSHighlightsV2.docx","url":"https://assets-eu.researchsquare.com/files/rs-8891770/v1/92520b423843f91b7e079a85.docx"},{"id":103505737,"identity":"d321369f-ef77-4595-8d3d-f801a170d241","added_by":"auto","created_at":"2026-02-26 13:32:50","extension":"png","order_by":3,"title":"","display":"","copyAsset":false,"role":"supplement","size":298259,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eGraphical Abstract\u003c/strong\u003e\u003c/p\u003e","description":"","filename":"GA.png","url":"https://assets-eu.researchsquare.com/files/rs-8891770/v1/f0a930955f072a41aaaf1fc2.png"}],"financialInterests":"No competing interests reported.","formattedTitle":"Phosphorus-arsenic interaction mitigates toxicity and accumulation of arsenic in rice grown in contaminated fields","fulltext":[{"header":"Introduction","content":"\u003cp\u003eThe presence of hazardous metal(loid) elements in soil and water is a growing global concern, posing serious threats to human and animal health and negatively impacting the well-being of plants and microorganisms (Majumdar et al., 2023; Tyagi et al., \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Unlike organic pollutants, metal(loid)s do not degrade and persist in soil for prolonged durations (Wang et al., \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Wang et al., \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Soil enriched with arsenic (As) is a significant source of contamination in the food chain and water supply, raising environmental and human health concerns due to the fact that As is a well-established carcinogen and mutagen (Chakrabarty \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Several soil parameters can influence As accumulation in rice, including total soil As concentration, redox status, poorly crystalline iron (Fe) (hydr)oxide concentration, plant available phosphorus (P), texture, pH, plant available silicon (Si), and sulfur (S) (Linam et al., \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Amaral et al., \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e \u003cp\u003ePhosphorus plays a critical role in regulating the uptake of As by plants through two primary processes. Arsenate [As(V)] serves as a phosphate analogue and is taken up by plants through a phosphate transporter system (Latowski et al., \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Therefore, As uptake and accumulation are known to be inversely influenced by P concentrations in the soil solution (Wu et al., \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Arsenic in soil exists in both inorganic and organic forms, including arsenite As(III), arsenate (As(V)), monomethylarsonic acid (MMA), and dimethylarsonic acid (DMA). Typically, As(V) is present as H\u003csub\u003e2\u003c/sub\u003eAsO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;3\u003c/sup\u003e or HAsO\u003csub\u003e2\u003c/sub\u003e\u003csup\u003e\u0026minus;4\u003c/sup\u003e, and As(III) as H\u003csub\u003e3\u003c/sub\u003eAsO\u003csub\u003e3\u003c/sub\u003e under circumneutral pH conditions (Huang et al., \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Seyfferth et al., \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). In the soil, phosphate competes for the same sorption sites as As(V) (Rivas-P\u0026eacute;rez et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). This competition facilitates the desorption of As into the soil solution, enabling its uptake by plants (Bakhat et al., 2017; Rahman and Hesegawa, 2011). Therefore, the impact of P on As mobilization depends on the soil's charge characteristics. In a hydroponic experiment, As-induced reductions in root growth and increased tissue As concentration were reversed by P treatment (Tu and Ma, \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2003\u003c/span\u003e). Therefore, P can effectively reduce As availability and root uptake in As-contaminated soils (Anwar et al., 2015). Cytoplasmic As(V) can disrupt metabolic processes involving phosphate, making it toxic to plants (Meharg and Hartley-Whitaker, \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). However, when internal phosphate concentration is high, it can lead to the down-regulation of phosphate transporters and reduce As uptake (Finnegan and Chen, \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Although As is not a redox-active metal, substantial evidence suggests that exposing plants to inorganic As generates reactive oxygen species (ROS), connected with valence changes of As from As(V) to As(III) in plants (Abbas et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Meharg and Hartley-Whitaker, \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2002\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe chemical nature of As(V) and phosphate, and their interactions in soil and plants, do indicate that P amendment can be a fruitful and practical strategy to regulate As levels in plants. However, it is important to exercise caution when amending soils with P to mitigate As uptake. In soils with low P concentrations and high As levels, increasing soil P levels can elevate As solubility, availability, uptake, and accumulation in rice grain due to P adsorption to soil exchange sites. Therefore, a balanced approach is required when using P to manage As contamination in the soil. In our previous work, two varieties differing in phosphorus use efficiency (PUE) were identified, and their interactions with As were studied in laboratory conditions (Shukla et al., \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Sai Kasturi (SK) was identified as a low PUE while DNA Sribala (DS) was identified as a high PUE variety. This study attempted to investigate phosphorus fertilization effects on arsenic uptake, oxidative stress responses, and crop productivity in rice crops under arsenic-contaminated soil conditions. Using two different varieties of rice defined by phosphorus utilization efficiency (PUE), research attempted to define genotype-related responses to varied phosphate supplies. The main goal was to find out if optimized phosphorus management, as a means to minimize arsenic content in rice grains, can still allow crop growth and production, providing an achievable and sustainable crop production practice to support food security in arsenic-affected areas.\u003c/p\u003e"},{"header":"Material and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eExperimental Site and Rice Cultivar Selection\u003c/h2\u003e \u003cp\u003eIn the study, As-contaminated fields were chosen to examine the effects of different P doses on particular rice types. The study was conducted in two different districts of Uttar Pradesh, India: Ghazipur (site 1) and Jaunpur (site 2) (Supplementary Figure \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e). These fields, which are located at 25.5084\u0026deg; N, 93.8925\u0026deg; E for Ghazipur and 25\u0026deg;34'22.1\" N, 82\u0026deg;58'32.0\" E for Jaunpur, were selected because of their comparable levels of contamination: Ghazipur (17.3 mg kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) and Jaunpur (16.5 mg kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e). Two selected rice cultivars were DNA sribala (DS) and Sai kasturi (SK), which were identified in our earlier lab study (Shukla et al., \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). DNA Sribala is a high PUE variety, responsive to phosphorus and tolerant to As stress under hydroponic conditions. Sai Kasturi is the low PUE variety and is As sensitive too (Shukla et al., \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). These rice varieties typically take between 125 to 135 days to reach maturity.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eField Layout, Irrigation, Experimental Design, and Fertilizer Application\u003c/h3\u003e\n\u003cp\u003eThe entire experiment was executed using a completely randomized block design. The field sites at two locations were divided into three blocks. The control block (normal As-contaminated soil at two sites) received a P dose of 60 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e. There were two experimental blocks: experimental blocks P1 (As-contaminated soil amended with 30 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e of P; half of the recommended P fertilizer) and P2 (As-contaminated soil amended with 120 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e of P; double the recommended P fertilizer). The seeds of both varieties were grown in nursery where soil-As concentration was less than 5 mg kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e. Two square shape (5*5 ft\u003csup\u003e2\u003c/sup\u003e) plots were prepared with submerged water condition (2\u0026ndash;3 cm). After 30 days after sowing (DAS) the rice plants with equal height were transferred into different experimental plots. Throughout the growth period (July to November), the control and experimental fields were kept at a constant water level of 3 to 6 cm. The ratio of (N:P: K) fertilizer applied per hectare was 120:60:40. (N:P: K) in control block, 120:30:40 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e in P1 and 120:120:40 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e in P2 with urea, superphosphate and muriate of potash being the respective sources. Nitrogen was applied in three stages: half during field preparation, and the remaining quarters were given during the early growth stage (tillering, 25DAS) and 50\u0026ndash;60 days after transplant (DAT). Weeding was managed manually four times throughout the entire experimental period, with the waterlogged conditions also contributing to weed control in the fields.\u003c/p\u003e\n\u003ch3\u003eSoil Sampling and Analysis\u003c/h3\u003e\n\u003cp\u003eArsenic-contaminated topsoil (0\u0026ndash;20 cm depth) was sampled at two different stages: During transplantation (30 DAS) and at maturity stage (135 DAS). Soil samples were obtained using PVC cores/pipes of 25 cm in length at each of these sampling points. The samples were subsequently air-dried, ground, passed through a 5 mm nylon sieve, and homogenized to create representative samples. The soil's physicochemical and biological properties, such as pH, electrical conductivity (EC), redox potential (ORP), and soil organic carbon (SOC), were analyzed using appropriate standard methods as detailed in Upadhyay et al. (\u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e\n\u003ch3\u003eHarvesting, Growth Measurement, and Yield Evaluation\u003c/h3\u003e\n\u003cp\u003ePlant sampling was done at two key time points throughout the cultivation period. The first sampling was done at 65 DAT, during the milking stage of cultivation. At this stage, the leaf samples were collected for the estimation of chlorophyll content and other biochemical assays. The second sampling was conducted at maturity (135 DAS). During this stage, root and shoot lengths were measured, spikes were separated, and fertile spikelet per spike were counted. Grains were separated, dried, and the weight of 1000 grains was determined to estimate the yield.\u003c/p\u003e\n\u003ch3\u003eMeasurement of Electrolytic Leakage and Lipid Peroxidation\u003c/h3\u003e\n\u003cp\u003eElectrolytic leakage (EL) was evaluated according to the protocol of Dionnisio-Sese and Tobita (1998). Fresh leaf samples (20 leaf discs) were submerged in 40 ml of deionized water, and after 10 minutes, electrical conductivity (EC1) was recorded. The solution was then subjected to controlled temperature baths at 50\u0026ndash;60\u0026deg;C for 25 minutes, and EC2 was measured. Lastly, the solution was boiled at 100\u0026deg;C for 10 minutes, and EC3 was recorded. Electrolytic leakage was calculated using the formula:\u003c/p\u003e \u003cp\u003eElectrolytic leakage (%) = \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:\\frac{(EC2-EC1)}{EC3}\\times\\:100\\)\u003c/span\u003e\u003c/span\u003e\u003c/p\u003e \u003cp\u003eLipid peroxidation was measured by estimating the production of malondialdehyde (MDA) using the protocol of Heath and Packer (\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e1968\u003c/span\u003e).\u003c/p\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eMeasurement of Pigments\u003c/h2\u003e \u003cp\u003ePigments, including total chlorophyll and carotenoids, were assessed from fresh leaf samples using chilled acetone extraction and spectrophotometric measurements. Chlorophyll and carotenoid concentrations were determined through established methodologies as outlined by Arnon (\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e1949\u003c/span\u003e) for chlorophyll and Duxbury and Yentsch (\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e1956\u003c/span\u003e) for carotenoids. Leaf samples weighing between 20\u0026ndash;25 mg were finely crushed and homogenized with 1 ml of 80% acetone, followed by centrifugation at 10,000 rpm for 15 minutes. The resulting supernatant was then employed for the analysis of chlorophyll a, chlorophyll b, and carotenoids, with absorbance readings taken at 480, 510, 645, and 663 nm using a UV-VIS spectrophotometer (Model-UV-1900, Shimadzu, India). Subsequent to the absorption measurements, the values were computed using specific calculations.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eProtein content determination and antioxidant enzyme assays\u003c/h3\u003e\n\u003cp\u003eProtein content determination utilized the Bradford (\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e1976\u003c/span\u003e) method. In this approach, a 1 ml sample, comprising 950 \u0026micro;l Bradford reagent and 50 \u0026micro;l enzyme extract, was meticulously mixed using a vortex. Following 10-minute incubation, the solution underwent a discernible transformation into a blue color, and the absorbance was measured at 650 nm. This method capitalizes on the distinctive color change at the specified wavelength for precise assessment of protein content. The activities of antioxidant enzymes superoxide dismutase (SOD), guaiacol peroxidase (GPX), ascorbate peroxidase (APX) and catalase (CAT) were assayed by following the methods of Beauchamp and Fridovich (\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1971\u003c/span\u003e), Hemeda and Klein (\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e1990\u003c/span\u003e), Nakano and Asada (1981) and Aebi (1974) as detailed previously (Srivastava et al., 2006).\u003c/p\u003e\n\u003ch3\u003eArsenic analysis\u003c/h3\u003e\n\u003cp\u003eThe plant samples were separated into different parts (root, flag leaves, grains). These parts were cleaned by gentle shaking and placed into beakers for oven-drying. Dried samples were collected, and grain samples were dehusked to get the edible grain samples and crushed into powder form. The samples were digested with HNO3 (70% Suprapur Merck; Darmstadt-Germany) in a heating test tube block digester at 180\u0026deg;C for 1 hour and then at 200 \u003csup\u003eo\u003c/sup\u003eC for 45\u0026ndash;60 minutes till the sample turned into a white crystalline appearance (Srivastava et al., \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Pathak et al., \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). After proper digestion, the final volume for each sample was maintained up to 10 ml with ultrapure (Type I) water (Milli-Q) and filtered with a 0.22 \u0026micro;m pore size filter paper. Finally, the P and As contents were measured through (HG-AAS). For hydride generation, analytical standard sodium borohydride (3%; Merck), sodium hydroxide (2.5%; Merck) and hydrochloric acid (6 M; Merck) were used. The precision of the analysis was checked by certified standard reference materials (SRMs) (NIST, USA), such as 1568a rice flour and 1573a tomato leaf, as per the details given in Khanam et al. (\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). The quality control was performed by triplicate analysis and by the measurement of the recovery of spiked digested samples. The recoveries varied from 95 to 98% for all the tested samples.\u003c/p\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eStatistical Analysis\u003c/h2\u003e \u003cp\u003eThe experiment was conducted using a Randomized Complete Block Design (RCBD) with three replications. Data were subjected to two-way analysis of variance (ANOVA) to evaluate the effects of location and treatment, as well as their interaction. When significant differences were detected, mean comparisons were performed using Tukey\u0026rsquo;s Honestly Significant Difference (HSD) test at a probability level of p\u0026thinsp;\u0026le;\u0026thinsp;0.05. Statistical analyses were carried out using GraphPad Prism.\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003eSoil physicochemical characteristics and soil enzymes\u003c/h2\u003e \u003cp\u003eThe soil samples were collected at two different times, and variations were observed in the blocks of both varieties under different P doses. The analysis of soil samples from both sites, represented in Supplementary Tables S1 and S2, revealed significant changes in various soil parameters. A decrease in pH was noticed from sampling at the vegetative stage to the heading phase, while EC was increased at both sites. There were some changes in ORP, soil organic carbon and enzymatic activities also. Soil As levels were also examined, showing changes from the initial concentrations for all treatments at both sites (supplementary Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e and S2). The level of soil As was generally decreased from vegetative to heading phase, and this decline was greater for SK blocks than for the DS blocks. In contrast to As, soil P levels were found to increase from the vegetative to the heading phase. These findings underscore the dynamic nature of soil properties and enzymatic activities in response to varying P doses, highlighting the importance of monitoring and understanding soil dynamics for sustainable agricultural practices.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003eMorphological differences in selected varieties\u003c/h2\u003e \u003cp\u003eThe influence of different levels of phosphorus fertilizer on the physical characteristics and yield of two varieties was noticeable. Both varieties showed a decline in their growth and yield when given only half the recommended P dose, while they displayed an increase with a high P dose. The SK variety experienced a more significant decrease. The DS variety showed better changes in root structure, with a 19% and 18% increase in length at Ghazipur and Jaunpur, respectively, with high P doses (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ea). In contrast, the SK variety showed a 14% and 3% increase at the same sites at high P dose. SK also experienced a greater decrease in shoot length, with reductions of 14% and 35% under low P doses at the respective sites, while DS showed a smaller decrease (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eb). Panicle length decreased notably for the SK variety under low P doses, while DS showed minimal reductions. The number of fertile spikelet decreased under low P doses, impacting grain weight and number per panicle. SK, being less efficient in P utilization, showed a more significant overall reduction. However, both varieties responded positively to increased P doses, leading to improved production and yield at both sites (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ea-f).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003eEffect on electrolytic leakage and MDA\u003c/h2\u003e \u003cp\u003eMalondialdehyde, an indicator of membrane damage, was used to assess the extent of stress under low P doses. MDA production in roots increased by 23% and 18% for SK, and 7% and 8% for DS at sites 1 and 2, respectively, under low P doses as compared to the control (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ea). In shoots, MDA increased by 17% and 11% for SK, and 6% and 2% for DS at the respective sites. However, at high P doses at both sites, either a slight increase or no significant change in MDA was noticed in both SK and DS. At P2 in Ghazipur, shoots of both SK and DS showed a significant decline in MDA than the control (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ea-b). Hence, high P dose proved to have an ameliorative effect.\u003c/p\u003e \u003cp\u003ePlants subjected to a reduced P fertilizer dose also exhibited elevated electrolytic leakage (EL) in both roots and shoots, indicating heightened stress levels. Root EL increased by 27% and 8% for SK, and 10% and 2% for DS at sites 1 and 2, respectively, compared to the control dose. Similarly, shoot EL increased by 23% and 19% for SK, and 5% and 14% for DS at sites 1 and 2, respectively (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ec-d). The rise in EL suggests compromised membrane integrity in response to nutrient stress. However, when plants received high P fertilizer supply, double the recommended dose, stress levels were reduced in both varieties compared to the half P and normal P doses (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ec-d).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec16\" class=\"Section2\"\u003e \u003ch2\u003eEffect on photosynthetic pigments\u003c/h2\u003e \u003cp\u003ePhosphorus, as a vital nutrient, plays a crucial role in the process of photosynthesis within plants. The deficiency of this essential nutrient can significantly impede the photosynthetic process. In our experiment, the supplementation of a low P dose resulted in a notable decrease in pigment levels, with a more pronounced effect being in the SK variety. The reduction in pigment content was statistically significant, revealing a decrease in chlorophyll a (chl a) by 29% and 27% for SK, and 14% and 18% for DS at sites 1 and 2, respectively. Chlorophyll b (chl b) exhibited reductions of 18% and 13% for SK, and 8% and 10% for DS at the respective sites, while carotenoid content decreased by 35% and 32% for SK, and 21% and 14% for DS at sites 1 and 2, respectively, compared to the control (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ea-c). Conversely, under conditions of high P dose, an increase in chl a, chl b, and carotenoid content was observed in comparison to the control. The increment in chl a was evident in both varieties, with a rise of 10% and 15% for SK, and 13% and 18% for DS at sites 1 and 2, respectively. Chl b showed increases of 30% and 27% for SK, and 34% and 33% for DS at the respective sites (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ea-c). These findings underscore the pivotal role of phosphorus in sustaining optimal pigment production, ultimately influencing the efficiency of the photosynthetic process in plants.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec17\" class=\"Section2\"\u003e \u003ch2\u003eResponses of antioxidant enzymes (SOD, APX, CAT and GPX)\u003c/h2\u003e \u003cp\u003eSuperoxide dismutase (SOD) activity was assessed in the roots and shoots of both selected plant varieties, revealing noteworthy variations in response to phosphorus (P) fertilizer levels. SOD activity exhibited a reduction under a low P dose. The reduction in SOD activity under low P doses was 11% and 24% for SK, and 4% and 10% for DS at sites 1 and 2, respectively. Similarly, in the shoots, a reduction in SOD activity was noted, which was 22% and 31% for SK, and 13% and 18% for DS at sites 1 and 2, respectively, compared to the control. Conversely, when plants received an adequate supply of P fertilizer, an increase in SOD activity was observed. In the roots, SOD activity rose by 9% and 36% for SK, and 9% and 25% for DS at sites 1 and 2, respectively. In the shoots, the increase in SOD activity was recorded as 17% and 16% for SK, and 5% and 8% for DS at sites 1 and 2, respectively, compared to the control dose. These findings highlight the responsive nature of SOD activity to varying P fertilizer levels (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ea-b).\u003c/p\u003e \u003cp\u003eA consistent trend was observed for APX activity, mirroring the patterns seen with SOD, in response to varying P fertilizer levels. APX activity in the roots and shoots exhibited a decrease when compared to the control dose for both varieties, with reductions being greater for SK than for DS at both sites (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ec-d). In contrast, under higher P doses, APX activity increased for both varieties at both sites. This rise in APX activity suggested a positive response to an increased supply of phosphorus, indicating the potential role of phosphorus in enhancing the antioxidant defense mechanisms mediated by APX.\u003c/p\u003e \u003cp\u003eThe activity of the CAT enzyme showed a reduction in the roots under a half P dose, with decreases of 25% and 8% for SK, and 20% and 12% for DS at sites 1 and 2, respectively. In shoots, CAT activity declined by 20% and 29% for SK, and 5% and 13% for DS at sites 1 and 2, respectively (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ea-b). Under a double P dose, CAT activity increased by 8% at site 1 and 6% at site 2 for DS, while for SK, there was an exceptional decrease of 6% at site 1 and an increase of 15% at site 2 (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ea-b). The decline in CAT activity was notable under a half P dose compared to the control, further emphasizing the impact of P availability on antioxidant enzyme activity in plants.\u003c/p\u003e \u003cp\u003eGPX activity in the roots exhibited a decline under a half phosphorus dose, with reductions of 30% and 13% for SK, and 1% and 3% for DS at sites 1 and 2, respectively. Similarly, in the shoots, GPX activity decreased by 13% for SK at site 1 and a significant 35% at site 2. For DS, there was a 10% increase at site 1, but a reduction of 6% at site 2 (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ec-d). Conversely, under double P dose, both root and shoot GPX expression increased for both varieties at both experimental sites, suggesting a positive response to higher P levels.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec18\" class=\"Section2\"\u003e \u003ch2\u003ePhosphorus and arsenic accumulation pattern in rice\u003c/h2\u003e \u003cp\u003eThe study revealed notable fluctuations in P accumulation patterns in response to varying doses of P fertilizer at two distinct sites, Ghazipur and Jaunpur. Alterations in P concentrations were observed across different plant parts, including the root, shoot, and grains, highlighting the responsiveness of these components to changes in P fertilizer dosage. The SK variety exhibited a significant reduction in P concentration; for example, a 17% decrease in root P accumulation under half-dose conditions at the Ghazipur site. Conversely, the DS variety displayed a non-significant reduction of only 3% in root P accumulation under similar conditions. Upon application of a double dose of P fertilizer, root P accumulation increased by 15% for the DS variety and 8% for the SK variety at site 1, with corresponding increases of 3% and 10% being noticed at site 2 (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003ea-c).\u003c/p\u003e \u003cp\u003eSignificant differences were observed in shoot and grain P accumulation for both varieties under varying P fertilizer doses. Under half-dose conditions, the SK variety experienced a substantial decrease of 21% and 20% in shoot P accumulation at the Ghazipur and Jaunpur sites, respectively, while the DS variety exhibited a negligible reduction. In terms of grain P accumulation, the DS variety demonstrated higher accumulation in both control and double P dose conditions, with negligible reduction under half-dose conditions. In contrast, the SK variety exhibited a noticeable reduction of 18\u0026ndash;20% in P accumulation for both sites under similar conditions (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003ea-c), highlighting varietal disparities in the response to P fertilizer doses across different plant parts.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eArsenic concentration within both the vegetative parts and grains exhibited sensitivity to variations in P fertilizer dosage. Under a low P dose, there was an elevation in As accumulation, whereas the addition of a double P dose resulted in a slight decrease. Notably, the accumulation of As was consistently higher in SK tissues and grains across all conditions, with an enhanced accumulation observed particularly under low P doses. In SK grains, the As concentration in control conditions was recorded at (0.58\u0026ndash;0.6 mg kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e). Under half the standard P dose, a significant increase to (0.69\u0026ndash;0.75 mg kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) was observed, indicating about a 20% rise at both contaminated sites. Conversely, applying a double P dose led to a reduction in As accumulation by 9\u0026ndash;11% at both sites (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003ea-c). In the case of the DS variety, the tissue and grain arsenic accumulation demonstrated resilience to lower P doses, with minimal effects. However, an application of higher P doses resulted in a significant reduction of As accumulation (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003ea-c). These findings underscore the differential response of arsenic accumulation in vegetative tissues and grains of SK and DS varieties to varying P fertilizer levels, offering valuable insights into the intricate interactions between phosphorus and arsenic dynamics in agricultural systems.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec19\" class=\"Section2\"\u003e \u003ch2\u003eRice yield of two cultivars grown in different fields under different P fertilizer doses\u003c/h2\u003e \u003cp\u003eThe assessment of rice yield, conducted within 100 ft\u003csup\u003e2\u003c/sup\u003e plots, revealed a range spanning from 5683 to 6496 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, as delineated in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e. A significant reduction in yield was seen under conditions of low P dosage. Specifically, at the Jaunpur site, the SK variety exhibited a substantial 31% decline, with yield plummeting from 3637 to 2492 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e. Conversely, the DS variety demonstrated a lesser reduction of 15% under analogous conditions. Parallel observations were made at the Ghazipur site, where the SK and DS varieties experienced reductions of 29% and 12%, respectively under low P (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Under high P, an increase in yield was observed for both varieties in both sites; the increase in yield ranged from 12\u0026ndash;14% for SK and 17\u0026ndash;20% for DS (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eYield parameters of rice varieties.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"6\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c4\" namest=\"c3\"\u003e \u003cp\u003eSK Variety\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e \u003cp\u003eDS Variety\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eYield (kg/100ft\u003csup\u003e2\u003c/sup\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eYield (kg/h)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eYield (kg/100ft\u003csup\u003e2\u003c/sup\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eYield (kg/h)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003e\u003cb\u003eJaunpur\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eControl\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3.38\u0026thinsp;\u0026plusmn;\u0026thinsp;0.31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3637\u0026thinsp;\u0026plusmn;\u0026thinsp;329.71\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3.88\u0026thinsp;\u0026plusmn;\u0026thinsp;0.22\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e4180\u0026thinsp;\u0026plusmn;\u0026thinsp;235.78\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eP1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.32\u0026thinsp;\u0026plusmn;\u0026thinsp;0.21\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2492\u0026thinsp;\u0026plusmn;\u0026thinsp;230.37\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3.29\u0026thinsp;\u0026plusmn;\u0026thinsp;0.25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e3537\u0026thinsp;\u0026plusmn;\u0026thinsp;268.72\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eP2\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3.86\u0026thinsp;\u0026plusmn;\u0026thinsp;0.23\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e4157\u0026thinsp;\u0026plusmn;\u0026thinsp;250.30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e4.65\u0026thinsp;\u0026plusmn;\u0026thinsp;0.24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e5007\u0026thinsp;\u0026plusmn;\u0026thinsp;255.16\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003e\u003cb\u003eGhazipur\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eControl\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.95\u0026thinsp;\u0026plusmn;\u0026thinsp;0.13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3179\u0026thinsp;\u0026plusmn;\u0026thinsp;141.06\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3.38\u0026thinsp;\u0026plusmn;\u0026thinsp;0.42\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e3641\u0026thinsp;\u0026plusmn;\u0026thinsp;456.77\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eP1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.08\u0026thinsp;\u0026plusmn;\u0026thinsp;0.16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2241\u0026thinsp;\u0026plusmn;\u0026thinsp;177.11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.99\u0026thinsp;\u0026plusmn;\u0026thinsp;0.41\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e3221\u0026thinsp;\u0026plusmn;\u0026thinsp;445.21\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eP2\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3.31\u0026thinsp;\u0026plusmn;\u0026thinsp;0.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3568\u0026thinsp;\u0026plusmn;\u0026thinsp;309.77\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3.94\u0026thinsp;\u0026plusmn;\u0026thinsp;0.46\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e4238\u0026thinsp;\u0026plusmn;\u0026thinsp;496.93\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003eRice (\u003cem\u003eOryza sativa\u003c/em\u003e L.), an essential food item for over half of the global population, is grown in over 100 countries, of which close to 90% of its total production is focused on the continent of Asia. Owing to its high calorific value, economic affordability, bland flavour, high iron content, and relatively low potential for allergenicity, rice forms an ideal constituent of the diet, particularly of children (Hoang et al., 2022). Arsenic (As) toxicity offers a two-sided threat by posing a threat not only towards the production of agriculture globally but also by being an effective human carcinogen. It is therefore essential that the problem be dealt with by improving crop sustainability as well as public health (Banerjee et al., \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe results of the present study propose that the DS variety exhibited superior performance in response to varying P fertilizer doses compared to the SK variety. Several key factors highlight the resilience and adaptability of the DS variety. Firstly, in terms of morphological attributes, the DS variety demonstrated better root and shoot growth under low P conditions, with a substantial increase in root length compared to SK (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ea-b). This superior root development may confer greater tolerance to nutrient stress, enabling DS plants to access and absorb nutrients more efficiently, ultimately contributing to enhanced overall growth even when the P is in scarcity. The evolution of higher plants has led to the development of various strategies for acquiring nutrients from the soil, especially for less available elements like P (Shen et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). One notable adaptation involves long root hairs at high densities in the roots of \u003cem\u003eArabidopsis\u003c/em\u003e plants. This adaptation allows plants to effectively explore a larger volume of soil and capture more nutrients (Lynch \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Thus, amended root growth is connected to better P nutrition of plants.\u003c/p\u003e \u003cp\u003eAdditionally, the DS variety unveiled more favourable responses in terms of shoot length, panicle length, and fertile spikelet development under varying P doses. The increased shoot and panicle lengths (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ec-f) suggest a greater above-ground growth, which is crucial for maximizing photosynthetic potential and overall biomass production. Moreover, the higher number of fertile spikelets indicates better reproductive success, contributing to potential increases in yield (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e) (Pretini et al., \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Diarra et al., \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e1985\u003c/span\u003e). Plant breeders often focus on optimized panicle structures for developing rice varieties with enhanced yield potential (Sabar et al., \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Previous studies on soybean, the effect on growth and yield was highly significant (α\u0026thinsp;\u0026lt;\u0026thinsp;0.05) with an increase in root/shoot dry and fresh weight in plants when a mixture of \u003cem\u003eBradyrhizobium\u003c/em\u003e inoculums along with phosphorus was used (Rabbani et al., \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Mirriam et al., \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Soil application of phosphorus resulted in a noteworthy rise in both yield and growth index in maize (Khaleeq et al., 2023). When initial phosphorus deficiency symptoms manifested 25 days after sowing in wheat, the application of higher doses of ammonium phosphate via foliar spray demonstrated the most significant reduction in phosphorus deficiency and led to the highest yields (Haloi, \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e1980\u003c/span\u003e). Thus, previous reports also support the contemporary idea. Figure\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003e summarizes the mechanism by which excess P lessens As toxicity in rice grown in As contaminated soil.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eIn the present study the morphological changes were conveyed by alterations in physiological processes. The grade of cell membrane damage (lipid peroxidation of membrane) and ROS buildup was determined by MDA and H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e quantification, and their pattern was consistent with cell death. Excessive production of ROS upon arsenic stress led to oxidation of cellular as well as organelle membrane lipids and MDA production. ROS triggers membrane lipid peroxidation by withdrawing an electron from the hydrogen atom of the fatty acyl chain of PUFA (polyunsaturated fatty acid), which makes the membrane rigid, leaky, and damaged membrane proteins (Abbas et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Disrupted membranes result in leakage of crucial components such as electrolytes from the cells, eventually leading to cell death (Salam et al., \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). In the present work, MDA accumulation reflected considerable differences between varying plant types as well as different levels of P fertilization. At low P availability, roots and shoots of SK reflected a noticeable elevation of MDA concentrations compared with DS, which shows that SK is rather more susceptible to oxidative damage. The higher MDA concentrations linked with P deficiency confirm increased production of reactive oxygen species (ROS) and lower membrane stability. By contrast, DS reflected relatively lower MDA concentrations that suggest higher inherent tolerance as well as more effective management of oxidative stress when faced with P limitations (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ea-d). Such genotypic differences in oxidative damage have been thoroughly reported for cereal species treated with nutrient stress regimes. Notably, at high P treatment doses, MDA concentrations either stayed the same or decreased, specifically at P2 in Ghazipur for shoots, wherein SK and DS significantly lowered from controls. This outcome indicates the beneficial action of sufficient P availability toward sustaining cellular redox balance as well as towards lowering membrane lipid peroxidation. Sufficient phosphorus is well documented for increased energy metabolism, enhanced nucleic acid and protein synthesis, and enhanced photosynthetic potential that mitigates ROS accumulation. This idea also finds corroboration from Al-Qahtani et al. (2024). Overall, these results underscore a bivalent role of phosphorus hereafter: deficiency of phosphorus causes oxidative stress as indicated by increased MDA accumulation, while sufficient or higher P supply mitigates oxidative membrane damage by enhancing enzymatic antioxidant defense.\u003c/p\u003e \u003cp\u003eAs per reports, photosynthetic activity is inhibited by arsenic tremendously due to the shifting that occurs in light-harvesting complexes that involve pigment losses as well as chlorophyll production. The Mg-ion of chlorophyll is replaced by As, disrupting photosynthesis (Banerjee et al., \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Figure\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e clearly depicts that reduced P doses hampered pigment production, leading to a substantial decrease in chlorophyll a, chlorophyll b, and carotenoid levels, with the SK variety experiencing more pronounced reductions than DS due to As toxicity. However, an adequate P supply mitigated these effects, resulting in increased pigment production by upregulating essential nutrient uptake.\u003c/p\u003e \u003cp\u003eIt is universally accepted that ROS hampers the overall health and productivity of plants. Superoxide dismutase (SOD), alongside glutathione peroxidase (GPX), is one of the key enzymatic antioxidants that scavenge ROS and cause subsequent alleviation of oxidative stress (Ighodaro and Akinloye, 2017). The results (Figs.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e and \u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e) clearly demonstrate that the antioxidant enzymatic activities of rice are strongly regulated by the availability of phosphorus (P). Consistent decline of SOD, APX, CAT, and GPX activities by half P rates indicates that the plant ROS-scavenging system is influenced by nutrient limitation, thereby enhancing susceptibility to oxidative stress. However, adequate P supply overcame and, for most instances, enhanced such enzymatic activities, indicative of adequate P fertilization sustaining redox homeostasis as well as stress resistance (Khan et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Among the enzymes studied, SOD and APX possessed the highest consistent positive response against higher P levels, which indicates their central role in the first defense against oxidative stress (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ea-c). CAT and GPX were more variable, particularly by site and variety, which reflects variation in their regulatory mechanisms. SK also responded more to P variation than did DS, which indicates genotypic variation of antioxidant controls (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ea-d). The results are in line with Mishra et al. (\u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), which showed similar evidence. In summary, varietal variation indicates potential for P-efficient genotypes possessing enhanced resistance against nutrient stress and provides useful insights towards sustainable nutrient use in rice production.\u003c/p\u003e \u003cp\u003eSome other studies showed that elevated P level has an ameliorative effect against various HM stress and influence plant growth and development. Arshad et al., (\u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) reported the provision of P enhanced shoot biomass, leaf area, photosynthetic pigments, and mineral nutrient levels, while decreasing concentrations of cadmium (Cd) and hydrogen peroxide (H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e) in wheat shoots. The application of P also improved antioxidant enzyme activities and gas exchange characteristics, serving as a crucial mechanism for enhancing Cd tolerance in wheat. A study conducted by Tairo et al. (2013) demonstrated that the addition of P led to a significant increase in growth attributes of soybean, such as the leaf area index, improved efficiency in photosynthesis, and promoted green pigment formation.\u003c/p\u003e \u003cp\u003eIn addition to biochemical and morphological changes, the major finding of the present study was the decline in As content of plant tissues, including grains, along with an enhancement of P content in high P supply. The research disclosed considerable varietal variation in P accumulation of different plant organs in response to different fertilizations. The SK variety was very sensitive to phosphate starvation, as evidenced by considerable decreases in P concentrations of root, shoot, and grains when they were supplied from half-doses during both locations of experiments. In contrast, the DS variety registered marginal decreases in root and shoot P contents while sustaining grain P accumulation, suggesting high tolerance to restricted phosphate supplies. Under double-dose fertilization, both genotypes registered greater P accrual, but DS recorded relatively larger increments in root and grain P compared with SK, suggesting greater utilization and acquisition efficiency. DS tended to maintain consistent P allocation across the root, shoot, and grain, while there was strong dependence on high external P application in SK. The study results (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e) showed that DS has improved adaptability to variable phosphorus availabilities, hence making it more suitable for farming on phosphorus-deficient soils. On the contrary, SK might require fertilizer management schedules tailor-made to sustain yield levels. The reduction in As content in roots, shoots, and grains could be attributed to the competition between As and P for the same transporters and binding sites. Arsenic enters rapidly across the root cells through phosphate transporters (PHTs) are characterized through their sequence homology with inorganic phosphate (Shukla et al., \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Banerjee et al., \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eHence, optimizing P nutrition to rice plants appears to be a viable and practical approach to mitigate As toxicity and reduce its accumulation in grains. Importantly, the reduction in As accumulation also resulted in higher yields of rice in high P supply. Hence, optimal P nutrition is a requisite of the plant that, when fulfilled, makes the plant grow and produce better. The present study also highlighted the importance of genetic variability in plant responses to nutrient stress (Basavaraj et al., \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). The study emphasizes the importance of selecting crop varieties with inherent traits that contribute to better nutrient utilization and stress tolerance. In conclusion, the findings of this study, coupled with relevant references, provide substantial evidence supporting the superiority of the DS variety under varying phosphorus conditions. This underscores the significance of genetic diversity in crop breeding programs and the potential for selecting varieties that can thrive under suboptimal nutrient conditions, contributing to sustainable and resilient agriculture.\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eThis study clearly establishes that phosphorus nutrition plays an overriding role in regulating arsenic dynamics, stress physiology, and yield performance in arsenic-contaminated soil-grown rice. Out of two cultivars tested, which one exhibited high and low phosphorus use efficiency (PUE), respectively, the high PUE variety DNA Sribala (DS) showed, on average, higher tolerance to nutrients as well as arsenic toxicity than the low PUE variety Sai Kasturi (SK). The increased tolerance exhibited by DS was expressed through greater root and shoot growth, higher panicle length, and higher number of fertile spikelets, and these, in totality, resulted in more stable and reproducible yield outcomes, even under low phosphorus conditions.\u003c/p\u003e \u003cp\u003ePhysiological and biochemical studies also showed that DS retained higher contents of photosynthetic pigments and possessed a better antioxidant defense mechanism, reflected in increased SOD, APX, GPX, and CAT activities under increased P supply. These adjustments effectively controlled oxidative damage, as reflected in decreased production of MDA and electrolyte leakage. Notably, P supplementation not only alleviated stress-caused damage but also substantially diminished As deposition in vegetative organs and grains, especially at elevated P doses. Though SK showed increased susceptibility to both low P and arsenic toxicity, DS showed an even and stable nutrient uptake profile, supporting the importance of P-efficient genotypes in contaminated agroecosystems.\u003c/p\u003e \u003cp\u003eIn short, this study shows double benefits: reducing arsenic seepage into the food supply and maximizing rice production. These results underscore the importance of precision nutrient management and judicious varietal selection as an environmentally sustainable agricultural practice to safeguard vulnerable areas threatened by arsenic. However, further studies are necessary to confirm these results under various soils and agro-climatic regimes to understand P-As interaction. Molecular research determine genes responsible for phosphorus use efficiency and arsenic tolerance would support breeding programs to develop resilient varieties. Integrating P-efficient varieties with superior nutrient management practices involving biofertilizers or nano fertilizers would also portray increased productivity and lower arsenic uptake. These measures, in conjunction with location-specific management guidelines, will be essential to provide crop safety and sustainable rice production in arsenic-contaminated areas.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors are thankful to IESD-BHU, Varanasi, India, for the lab facility. SS is thankful to IOE-BHU (No.-6031) for financial support. KS is thankful to CSIR for Senior Research Fellowship (09/013(0818)/2018-EMR-I).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eKS executed all the experiments, formal analysis, and data curation, and prepared the draft the SKP helped in formal analysis and writing the original draft. SB helped in writing the original draft. SS conceptualized and supervised the study and finalized the draft for submission.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAvailability of data and materials\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe data that support the findings of this study are available from the corresponding author upon reasonable request.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eDeclaration of interests\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare no conflict of interest.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAbbas, G., Murtaza, B., Bibi, I., Shahid, M., Niazi, N. K., Khan, M. I., \u0026hellip; Natasha. (2018). 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Rhizospheric nano-remediation salvages arsenic genotoxicity: Zinc-oxide nanoparticles articulate better oxidative stress management, reduce arsenic uptake, and increase yield in Pisum sativum (L.). \u003cem\u003eScience of the Total Environment\u003c/em\u003e, \u003cem\u003e913\u003c/em\u003e, p.169493.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMishra, R.K., Mishra, G., Singh, R., Parihar, P., Kumar, J., Srivastava, P.K. and Prasad, S.M., 2022. Managing arsenic (V) toxicity by phosphate supplementation in rice seedlings: modulations in AsA-GSH cycle and other antioxidant enzymes. \u003cem\u003eEnvironmental Science and Pollution Research\u003c/em\u003e, \u003cem\u003e29\u003c/em\u003e(10), pp.14418\u0026ndash;14429.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAl-Qahtani, W.H., Abdel-Maksoud, M.A., Saleh, I.A., Zomot, N., Almunqedhi, B.M., Kataya, A.R., Fahad, S. and Ali, S., 2025. Effects of Phosphorus Sources on Arsenic Stress Mitigation in Wheat via Proline and Antioxidant Pathways. \u003cem\u003eJournal of Plant Growth Regulation\u003c/em\u003e, \u003cem\u003e44\u003c/em\u003e(2), pp.821\u0026ndash;838.\u003c/span\u003e\u003c/li\u003e\u003c/ol\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":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"plant-cell-reports","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"pcre","sideBox":"Learn more about [Plant Cell Reports](https://www.springer.com/journal/299)","snPcode":"299","submissionUrl":"https://submission.nature.com/new-submission/299/3","title":"Plant Cell Reports","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Antioxidant enzymes, Crop productivity, Food safety, Phosphorus accumulation, Phosphorus use efficiency","lastPublishedDoi":"10.21203/rs.3.rs-8891770/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8891770/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eArsenic (As) contamination in rice is a serious threat to food security worldwide. Phosphorus (P) plays a crucial role in modulating As uptake in rice plants. Elevated P content in the soil may hinder the uptake of As by competing with the transporter system and altering rhizosphere chemistry.\u003cstrong\u003e \u003c/strong\u003eThis study aimed to investigate the effect of varied P fertilizer levels on rice (\u003cem\u003eOryza sativa \u003c/em\u003eL.) plants grown in As-contaminated soil in terms of growth and As accumulation.\u003cstrong\u003e \u003c/strong\u003eThe experiment included recommended (60 kg h\u003csup\u003e-1\u003c/sup\u003e), half (30 kg h\u003csup\u003e-1\u003c/sup\u003e), and double (120 kg h\u003csup\u003e-1\u003c/sup\u003e) doses of diammonium phosphate (DAP). Two contrasting varieties of rice were used that differed in phosphorus use efficiency (PUE): P-efficient DNA Sribala (DS) and P-inefficient Sai Kasturi (SK). SK suffered significant reductions in shoot and panicle lengths under low P doses (up to 35% and 18%, respectively), whereas DS showed non-significant effects. Arsenic accumulation in grains increased by 20% under low P doses for SK but decreased by 9-11% with higher P doses. In contrast, DS demonstrated tolerance to low phosphorus concentrations and no significant change in As accumulation was observed. In addition, the yield of rice was also increased in high P doses, which was 14-23% at two sites for SK, while 17-20% for DS.\u003cstrong\u003e \u003c/strong\u003eThese findings underscore the critical role of P management in mitigating As toxicity and optimizing rice productivity in contaminated environments.\u003c/p\u003e","manuscriptTitle":"Phosphorus-arsenic interaction mitigates toxicity and accumulation of arsenic in rice grown in contaminated fields","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-02-23 16:21:06","doi":"10.21203/rs.3.rs-8891770/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2026-02-27T06:23:20+00:00","index":"","fulltext":""},{"type":"reviewerAgreed","content":"292550103601737518600497835623848226525","date":"2026-02-25T05:41:17+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-02-24T15:22:04+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-02-24T13:48:18+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-02-22T06:02:34+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"78550421142632627823274397505584556456","date":"2026-02-21T09:18:25+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"267607071186305700901922026088167224627","date":"2026-02-21T07:53:45+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"50549850707946826718965029744294677800","date":"2026-02-21T06:53:34+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"146576000382317139219989584882948592577","date":"2026-02-19T16:26:45+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"322436836788309840797606191278993339664","date":"2026-02-19T10:09:53+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"334250726738450012118705957088770061923","date":"2026-02-19T06:56:27+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"281916791030223130644488224756777745799","date":"2026-02-19T06:42:15+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2026-02-19T06:21:34+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2026-02-18T06:41:52+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2026-02-18T05:23:11+00:00","index":"","fulltext":""},{"type":"submitted","content":"Plant Cell Reports","date":"2026-02-16T09:23:18+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"plant-cell-reports","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"pcre","sideBox":"Learn more about [Plant Cell Reports](https://www.springer.com/journal/299)","snPcode":"299","submissionUrl":"https://submission.nature.com/new-submission/299/3","title":"Plant Cell Reports","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"238d3cc8-6dda-4990-9210-0afc3cdac016","owner":[],"postedDate":"February 23rd, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-04-10T05:40:48+00:00","versionOfRecord":[],"versionCreatedAt":"2026-02-23 16:21:06","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8891770","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8891770","identity":"rs-8891770","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}