Investigation of Phytochemical Characteristics of Quinoa in an Intercropping Production System with Maize in Response to Mycorrhiza and Phosphorus Fertilizer

Investigation of Phytochemical Characteristics of Quinoa in an Intercropping Production System with Maize in Response to Mycorrhiza and Phosphorus Fertilizer | 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 Article Investigation of Phytochemical Characteristics of Quinoa in an Intercropping Production System with Maize in Response to Mycorrhiza and Phosphorus Fertilizer Tayebe Rostami, Hamid Abbasdokht, Hassan Makarian, Manoochehr Gholipoor, and 1 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-3930581/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Incorporating bio-fertilizers, such as arbuscular mycorrhizal fungi (AMF), into mixed cropping systems has emerged as an innovative and sustainable approach to enhance crop growth and promote sustainable production. The study was conducted in two distinct locations within a mixed cropping system of corn and quinoa. Phosphorus fertilizer was applied at three different levels 50, 100, and 150 kg/ha while also considering the presence or absence of mycorrhiza. Cropping ratios included individual cultivation of quinoa, maize, and intercrop ratios of 50: 50, 75: 25, and 25: 75 (maize: quinoa). The utilization of phosphorus fertilizer improved the symbiotic interaction between plants and arbuscular mycorrhizal fungi, resulting in an increased rate of inoculation by AMF. The 50: 50 intercropping ratio, combined with the application of AMF, resulted in the highest contents of flavonoids, total phenolic compounds, tannins, saponin, antioxidant activity, and phytic acid in quinoa. Our findings demonstrated that AMF exhibit specific roles in optimizing plant strategies for phosphorus uptake and the efficient utilization of phosphorus in the mixed cropping. Recommending the application of AMF in a 50:50 intercropping ratio could be proposed to farmers as a friendly method to attain favorable phytochemical characteristics, particularly when using 50 kg/ha of phosphorus fertilizer. Biological sciences/Chemical biology Biological sciences/Ecology Biological sciences/Physiology Mixed cropping Quinoa Arbuscular mycorrhizal fungi Phytochemical traits and Phosphorus fertilizer. Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Introduction The United Nations' 2030 Agenda for Sustainable Development emphasizes the importance of establishing sustainable food systems that can provide nutritious diets for a growing global population. This means prioritizing crops that guarantee improved efficiency in using natural resources and contribute to meeting the requirements of a healthy diet. In line with this objective, alternative crops, such as quinoa, have received significant attention [1]. Quinoa ( Chenopodium quinoa Willd.) is an annual crop that belongs to the Chenopodiaceae family. It is native to the Andean region in South America and is primarily cultivated in countries such as Peru, Bolivia, Ecuador, Argentina, Colombia, and Chile [2]. According to a report by the Food and Agriculture Organization (FAO), the quinoa cultivation area has expanded from 52,555 hectares in 1961 to 184,585 hectares in 2019. Similarly, quinoa production has risen from 32,435 tons in 1961 to 161,415 tons in 2019 [3]. Quinoa exhibits resilience to agroecological extremes, including variations in soil conditions, rainfall, temperature, and altitude. It can thrive at sea level up to 4500 m above sea level, demonstrating adaptability to frost, drought, and salinity [4]. In contemporary times, quinoa has garnered increased attention due to its exceptional adaptability to diverse environmental conditions, substantial nutritional content, and widespread preference as an ingredient in gluten-free and functional foods. Being gluten-free, it serves as a source of protein, fatty acids, tocopherol, phytosterols, phenolic compounds, and has a low glycemic index [5]. Moreover, quinoa stands out for its abundant natural antioxidants, including ascorbic acid, flavonoids, phenolic compounds, and phytosterols [6, 7]. In today's world, implementing multi-cropping systems has become crucial, particularly in developing countries and among smallholder farmers. This approach allows for the efficient utilization of land and resources, while also reducing the incidence of pests and diseases [8, 9]. Intercropping agricultural systems have demonstrated notable efficacy in boosting production, contingent on specific ecological conditions. Intercropping is particularly preferred in regions with constraints such as limited land availability, labor force, and mechanization, as it enables the optimal utilization of available resources compared to single-cropping systems [10]. Phosphorus is a critical element for the growth and development of plants, playing a vital role in crop production. However, phosphorus deficiency has emerged as a significant constraint on agricultural productivity [11]. Research conducted by Dang et al. [12] on mung bean ( Vigna radiata L.) demonstrated that the application of phosphorus fertilizer led to a notable increase in the activities of superoxide dismutase and peroxidase, by 20.9% and 43.7% respectively. Arbuscular mycorrhizal fungi (AMF) mycelium plays a pivotal role in boosting soil phosphatase and microbial activity, significantly influencing phosphorus availability. In symbiosis with plants, especially crops, AMF fosters root growth and facilitates the conversion of inorganic phosphorus to organic forms [13]. These fungi form symbiotic relationships with various crops, residing both on root surfaces and within the epidermis of root cells. Intercropping soybean and maize, along with AMF inoculation, has shown significant benefits in promoting the growth of both maize and soybean. Furthermore, this approach has led to increased concentrations of available nitrogen and phosphorus in the soil compared to monoculture systems. Additionally, studies have indicated a positive correlation between the availability of phosphorus fertilizer and plant biomass [14]. These findings underscore the importance of phosphorus management strategies in optimizing plant growth and maximizing agricultural productivity. In their study, Amani Machiani et al. [15] found that the application of AMF had a positive impact on the production of secondary metabolites. The researchers specifically investigated the intercropping of soybean and thyme, combined with AMF application. They observed that this intercropping approach led to improvements in both yield and the production of secondary metabolites in thyme [16]. These findings highlight the potential of AMF utilization and intercropping strategies in enhancing the synthesis of valuable secondary metabolites, offering promising avenues for improving crop productivity and quality. This study investigated the influence of mycorrhiza inoculation and phosphorus fertilizer application on the phytochemical traits of quinoa within an intercropping system, considering two different climatic conditions. It is noteworthy that the observed colonization of quinoa by mycorrhizal fungi stands as one of the first documented instances of such symbiotic relationships in Iran. The findings of this study contribute to our understanding of how mycorrhiza inoculation and phosphorus fertilizer application can influence the phytochemical composition of quinoa in different environmental conditions. Results Total Flavonoids Content In Table 3, the significant influence of various treatments on the phytochemical properties of quinoa is demonstrated. The highest flavonoid content (66.80 mg g -1 ) was observed when cultivating 100% quinoa with mycorrhiza inoculation and applying 50 kg ha -1 phosphorus at the Mayamey site. On the flip side, the Shahrood location exhibited the lowest flavonoid content (41.5 mg g -1 ) when adopting a planting combination of 25% quinoa + 75% maize, coupled with 100 kg ha -1 phosphorus application (without mycorrhiza utilization) (Figure 1). Treatments involving mycorrhiza application exhibited superior effectiveness when contrasted with those lacking mycorrhiza inoculation. The diminished flavonoid content in the Shahrood location, particularly in the cultivation blend of 25% quinoa and 75% corn, may be ascribed to the climatic and soil conditions prevalent in this area (Figure 1), and coupled with light shortage due to the shading of corn on quinoa. Furthermore, a positive connection was observed among flavonoids, total phenol, and tannin contents. This implies that the surge in secondary metabolites stems from improved nutrient absorption facilitated by mycorrhiza hyphae (Table 5). Total Phenolic Compound (TPC) The analysis of variance results reveals a significant impact of intercropping, phosphorus fertilization, and mycorrhiza treatment on the total phenolic compounds content (TPC) of quinoa at two distinct sites (Table 3). The highest TPC value (40.57 mg g -1 ) was detected at the Mayamey location with the combined utilization of 50 kg ha -1 phosphorus and mycorrhiza, as well as at the Shahrood site with mycorrhiza application and absence of phosphorus. Conversely, the lowest TPC value (29.5 mg g -1 ) was documented at the Shahrood location when applying 100 kg ha -1 phosphorus (Figure 2). Tannins content The influence of varying phosphorus fertilizer levels, geographical locations, and intercropping on tannin levels exhibited a notable significance at a 1% level. Furthermore, all main effects and the interactive effects of cropping ratio, locations, phosphorus fertilizer, and mycorrhizal application were significant at the 1% level (Table 3). Examination of mean comparisons unveiled that the peak tannin content (0.49 mg g -1 ) was identified in the intercropping of 50% quinoa + 50% maize, accompanied by the application of 50 kg ha -1 phosphorus and mycorrhiza, at the Mayamey location. On the contrary, the least tannin content (0.11 mg g -1 ) was documented under the intercropping of 25% quinoa + 75% maize in the Shahrood locale, in the absence of mycorrhiza utilization (Figure 3). Additionally, a notable positive correlation was observed between the tannin content and the antioxidant activity, alongside antioxidant enzymes (Table 5). Saponin content The saponin content was significantly influenced (p<0.001) by the three-way interactions involving cropping ratio, locations, and phosphorus fertilizer (Table 3). The results indicated a reduction in saponin content as the ratio of corn increased in the intercropping, evident in both the control group and the application of phosphorus (Figure 4). Additionally, a strong positive correlation at the 1% significance level was found between saponin content and the levels of tannins and flavonoids (Table 5). Phytic acid content The phytic acid content in quinoa was significantly impacted by all individual treatments (p<0.001). However, most interactive treatments did not show statistically significant effects on phytic acid content. The treatment with the highest phytic acid content (3.8%) was observed in the intercropping of 50% quinoa and 50% maize with a phosphorus application of 50 kg ha -1 (Table 3). Conversely, the lowest phytic acid content (2.68%) was found in the intercropping of 25% quinoa + 75% maize in combination with phosphorus application at 100 kg ha -1 in the Shahrood location (Figure 5). Antioxidant The cropping ratio, phosphorus fertilizer application, mycorrhiza inoculation, and locations had a significant impact on the antioxidant activity in quinoa (Table 3). Upon examining the interactive effects of phosphorus levels with mycorrhiza inoculation, it was found that the highest antioxidant activity (79.85%) was attained with the application of 50 kg ha -1 of phosphorus, mycorrhiza treatment, and a cropping ratio of 50% quinoa. Conversely, the lowest antioxidant activity (52.05%) was recorded in the intercropping of 25% quinoa ratio, phosphorus application of 100 kg ha -1 , and the absence of mycorrhiza (Figure 6). Additionally, a significant positive correlation emerged between antioxidant activity and the levels of total phenols, flavonoids, tannins, as well as antioxidant enzymes (CAT, POX, SOD) (Table 5). Antioxidant Enzymes (CAT, POX, SOD) The antioxidant enzyme contents were notably impacted by the cropping ratio, phosphorus application, and mycorrhiza inoculation (P≤0.05). Elevated levels of antioxidant enzymes were evident with mycorrhizal and phosphorus applications when compared to the control conditions (Table 3). In the Mayamey location, the utilization of mycorrhizal treatment and phosphorus application (50 kg ha -1 ) resulted in higher antioxidant enzyme contents than those observed in the Shahrood location. The highest antioxidant enzyme contents were witnessed with the application of phosphorus fertilizer at 50 kg ha -1 , while the treatment without phosphorus application and mycorrhiza inoculation exhibited the lowest antioxidant enzyme contents (Table 4). Colonization Percentage The various treatments, encompassing locations, mycorrhiza introduction, phosphorus usage, and intercropping ratio, exhibited a significant influence (p<0.001) on the colonization percentage in quinoa (Table 3). Through mean comparison analysis, it was evident that the highest colonization rate (44.6%) was registered in the intercropping consisting of 25% quinoa and 75% maize, along with the application of 50 kg ha -1 of phosphorus and the AMF application. Conversely, the lowest colonization rate (5%) was noted in the 100% quinoa cultivation system without phosphorus application (control) in the Mayamey location (Figure 7). A strong and noteworthy correlation was found between the levels of flavonoids and phenols, along with the antioxidant enzyme content (Table 5). In contrast, no significant correlation was identified between phytic acid content and the measured phytochemical properties, encompassing antioxidant levels and related enzymes. The colonization percentage exhibited a significant positive correlation with both flavonoid and phytic acid levels (P≤0.05). Moreover, no noteworthy correlation was identified between saponin content and the antioxidant enzymes content. Discussion In this research, plants subjected to mycorrhizal fungi exhibited an expanded root system, enhancing the availability of water and essential nutrients, notably nitrogen, phosphorus, and potassium, crucial for plant growth and development [17]. The mycorrhizal hyphae played a role in augmenting magnesium absorption, consequently elevating the levels of total chlorophyll and flavonoids in both quinoa and maize plants [18]. The synthesis of total chlorophyll is contingent upon the adequate absorption of minerals, particularly nitrogen and magnesium [19]. Phosphorus plays a crucial physiological role in the biosynthesis pathway of biochemical compositions, as it exhibits a synergistic effect with nitrogen. Our findings indicate that the utilization of AMF resulted in an augmentation of secondary metabolite levels in quinoa compared to the control treatment. This increase may be attributed to the enhanced biochemical compositions [18]. Secondary metabolites, such as flavonoids belonging to phenylpropanoids, play a crucial role in maintaining the structural and functional integrity of cells, thereby mitigating oxidative damage caused by ROS. This protective mechanism aids plants in coping with drought stress [20, 21]. AMF has been observed to significantly enhance anthocyanin and total flavonoid concentrations in Medicago truncatula leaves [22]. It's noteworthy that these mycorrhizal effects can, in part, be attributed to the improved phosphorus (P) content [23]. Xia et al. [21] demonstrated that mycorrhizae lead to the accumulation of root metabolites. However, the exact roles of these secondary metabolites induced by AM symbiosis in enhancing plant drought tolerance remain largely unknown. One of the well-known benefits of AM symbiosis to the host plant is the significant improvement in water and nutrient availability facilitated by AM. This, in turn, can explain the changes observed in carbohydrates and certain components of secondary metabolism, which are referred to as general mycorrhizal effects [24]. While the effects of mycorrhiza on plants are frequently considered to be highly specific to host plants and/or fungal species, there are common characteristics in the alterations of mycorrhiza-associated metabolites in plant roots [25]. In line with our findings, previous studies have reported that mycorrhizal associations often lead to increased levels of bioactive phenolic and lignin metabolites compared to plants not inoculated with AM [26, 27]. Furthermore, mycorrhizal symbiosis can induce plant-specific metabolic changes, particularly in stress conditions, which cannot be fully explained solely by nutrient effects [22]. Experimental data and meta-analysis have unveiled highly specific effects of mycorrhizal fungi, plant species, and/or their interactions on plant metabolism [17]. The growth, development, and metabolic activity of plants are contingent upon their photosynthetic capacity and ability to utilize soil nutrients [28]. By introducing phosphorus supplementation in conjunction with AMF, the levels of available nutrients can be modified, thereby leading to alterations in the metabolic environment of plants. This, in turn, facilitates the enhanced absorption and assimilation of CO 2 from the atmosphere. The presence of mycorrhizal fungi resulted in enhanced biochemical properties, including increased antioxidant enzymes activity, flavonoids content, and total phenol, achieved by reducing peroxidase activity [29]. The hyphae of arbuscular mycorrhizal fungi possess the capability to augment water and nutrient uptake by plants, thereby establishing a symbiotic relationship between plants and AMF [30]. Quinoa seeds are known for their abundance of bioactive compounds, which display potent antioxidant activity and contain natural antioxidant compounds [31]. The total phenolic compounds present in quinoa contribute to its antioxidant activity [32]. According to Nsimba et al. [33], the total phenol content of quinoa extract grown in Japan was measured to be 148.0 mg g -1 equivalent of tannic acid, which was higher compared to the total phenol content (94.3 mg g -1 ) of quinoa grown in Bolivia. The disparities in phenolic component content between these two quinoa ecotypes can be attributed to variations in environmental factors and agricultural conditions [32]. Similar findings were observed in our experiment. Our results indicated that in the Shahrood region, the highest phenolic component content was observed in the treatment involving mycorrhiza inoculation alone. However, in the Mayamey region, the application of mycorrhiza and 50 kg ha -1 of phosphorous resulted in the highest phenolic component content (Figure 3). The total phenolic component content in quinoa seeds varied between 16.8 and 59.7 mg/100 g, whereas the content of soluble phenolic compounds in the samples ranged from 7 to 61 mg/100 g [31].While saponins present in quinoa have been identified as anti-quality factors, isolated saponins also possess intriguing biological properties [34]. Our findings revealed that the saponins content decreased as the proportion of maize in mixed crops increased (Figure 3). The application of 100 kg ha -1 phosphorus significantly reduced the saponin levels compared to the control and 50 kg ha -1 phosphorus in the 75% maize proportion. It has been reported that saponins may exhibit nutritional or pharmacological value and their levels can vary under different conditions [35].Tannins, belonging to the group of polyphenols, can form complexes with other macromolecules, leading to various biological effects, both beneficial and undesirable. Polyphenols are natural substances that are widespread and commonly distributed in the plant kingdom [36]. Phenolic compounds, flavonoids, and saponins, which are products of secondary metabolism, exhibit crucial antioxidant activities that promote cardiovascular and cerebrovascular health. Additionally, they possess antiallergenic, antibacterial, anticarcinogenic, and anti-inflammatory effects [37]. Intercropping systems in agriculture have demonstrated significant effectiveness in enhancing production, contingent upon specific ecological conditions [38]. In a study conducted by Koca [39] on the forage yield and growth parameters of the maize-quinoa intercropping system, it was observed that the amount of maize dry matter increased in nearly all mixtures. Li and Cai [28] observed that enhancing the presence of AMF led to significant improvements in various maize traits, including biomass, chlorophyll content, plant height, and leaf area. AMF mycelium plays a crucial role in enhancing phosphatase activity and microbial activity in the soil. These fungi have a significant impact on the availability of phosphorus in the soil. Through their symbiotic relationship with plants, particularly crops, AMF promotes root growth and facilitates the conversion of inorganic phosphorus into organic forms [13]. AMF can be found in symbiosis with a wide range of crops, residing on the surface of roots and within the epidermis of root cells. The hyphae of mycorrhizal fungi are finer than root hairs, allowing them to penetrate soil micropores and transport certain minerals such as nitrates and phosphates to their host plant in exchange for carbohydrates [40]. This aligns with the findings of the current study. Consequently, plants associated with mycorrhizae generally exhibit greater drought tolerance compared to those without mycorrhizal associations [41]. Phenolic compounds, as secondary metabolites, play a role in detoxification, aiding in the neutralization of active oxygen species and free radicals before they can harm plant cells. The antioxidant potential of a plant is contingent upon the concentration of phenolic and tannin compounds. This study demonstrated an increase in the amount of phenolic and tannin compounds when AMF were utilized. Under stress conditions, mycorrhizal plants respond to oxidative stress by enhancing the synthesis of phenolic compounds. Previous research has also reported an increase in phenolic and tannin compounds facilitated by AMF in various plants, including violets [42], grapes [43], and Commiphora leptophloeos [44]. In our study, variations in the saponin content of quinoa were observed. As per the results, the saponin content decreased with an increase in the proportion of maize in mixed crops (Figure 3). The application of 100 kg ha -1 phosphorus significantly reduced the saponin content compared to the control and 50 kg ha -1 phosphorus in the 75% maize proportion. Additionally, when quinoa samples underwent different pearling processes, the saponin levels decreased [45]. These findings highlight the influence of cropping proportion, phosphorus application, and processing techniques on the saponin content of quinoa. Moreover, the quantities and composition of saponins in the quinoa plant were not consistently stable due to the continual removal of seed pericarps and bran in the harvesting process to render it edible, for instance, through pressure-cooking and toasting [45]. The total saponin content in quinoa grown ranged from 3.81 to 27.1 mg gr -1 in Washington State [46, 47], aligning with our findings. An increase in the nitrogen (N) uptake rate per root length has been shown to enhance saponin accumulation in roots, indicating a significant positive relationship between N uptake rate per root length and saponin content in taproots. Sufficient nitrogen fertilizer has the potential to enhance both root structure and nutrient uptake efficiency, subsequently promoting saponin synthesis [48]. Given the crucial role of nitrogen in saponin synthesis, the reduction in saponin content observed in high concentrations of phosphorus (100 kg ha -1 ) in this study may be attributed to a decrease in nitrogen absorption and, consequently, a decrease in saponin contents. Andrino et al. [49] demonstrated the presence of mono-, di-, and tricarboxylic low-molecular-weight organic acids in compartments containing orthophosphate (OP) or goethite-bound-orthophosphate (GOE-PA) and phytic acid (PA or GOE-PA), indicating the occurrence of reductive dissolution and ligand exchange/dissolution reactions. Additionally, hyphae grown in goethite loaded with OP and PA exhibited an elevated content of unsaturated lipids, suggesting increased membrane fluidity to maintain optimal hyphal functionality and facilitate phosphorus incorporation. In the presence of plants, AMF demonstrates extensive practical diversity and contributes to growth-promoting functions by producing various metabolites. These functions include the mineralization of phytate, the production of siderophores, the dissolution of phosphorus, and the production of low-molecular-weight organic acids [50]. Additionally, the mycelium of rhizobium plays a role in mobilizing phosphorus from acid phytic, as highlighted by Selvakumar et al. [51]. AMF contribute to phytate mineralization, leading to the transfer of phosphorus [52]. These studies collectively emphasize the significant contributions of AMF in promoting plant growth by facilitating the mineralization of phytate and enhancing phosphorus availability through various mechanisms. While previous studies have predominantly focused on the mechanisms of interaction between plants and beneficial microbes, such as bacteria and AMF, with an emphasis on model and crop plants [53, 25], the associated transcriptomic and metabolomic changes induced by AMF to alleviate effects of adverse environmental conditions in non-model plant species have not been fully elucidated. The photosynthetic capacity and ability to utilize soil nutrients are crucial factors that influence the growth, development, metabolic activities, and fatty acid composition of plants [28]. Zamani et al. [54] demonstrated that the amounts of phenolic compounds and fatty acid profiles in the intercropping of Lallemantia iberica and Cicer arietinum L. increased with the application of AMF. In the 50% intercropping pattern of quinoa with the application of AMF, the levels of phytic acid in quinoa increased. This can be attributed to the improved accessibility of nutrients facilitated by AMF, which in turn affects plant growth and photosynthesis. These changes in nutrient availability and metabolic processes can influence the production of fatty acid precursor compounds and the activity of enzymes like fatty acid synthase and acetyl-CoA carboxylase [55]. Furthermore, other studies have highlighted the positive effects of intercropping systems on plant nutrient availability, oil productivity, and oil quality. For instance, Rezaei Chiyaneh et al. [56] demonstrated that a cropping ratio of 50:50 in an intercrop of black cumin with fenugreek, using biofertilizer, resulted in the highest oil concentration in black cumin. This increase in oil concentration can be attributed to the enhanced nutrient uptake facilitated by the intercropping system. These findings emphasize the potential benefits of incorporating AMF and intercropping practices with biofertilizers in enhancing nutrient availability, oil production, and oil quality in various plant species. Rezapour et al. [57] demonstrated that the joint application of phosphorus and AMF resulted in an enhanced synthesis of fatty acids in groundnut. Additionally, the application of phosphorus fertilizer in a groundnut-corn intercropping system with a ratio of 1:2 improved groundnut oil content [57]. In a parallel study, it was found that the combined use of phosphorus chemical fertilizer and biological fertilizer increased safflower oil yield. These findings highlight the potential of using phosphorus fertilizers in conjunction with AMF and intercropping systems to enhance fatty acid synthesis and improve the oil content and yield of various oilseed crops like groundnut and safflower [58]. Oil synthesis, compared to other compounds studied in this research, requires a greater amount of energy. As a result, the utilization of mycorrhiza plays a significant role in enhancing the availability and absorption of nutrients by the plant root, providing the necessary energy for oil synthesis. This increased nutrient uptake facilitated by mycorrhiza ultimately leads to an augmentation in the production of oil. Furthermore, the energy required by plant processes is supplied in the form of high-energy ATP molecules. Phosphorus plays a crucial role in the formation of these energy-rich molecules. Therefore, the availability of phosphorus in the plant system contributes to the production of more high-energy ATP molecules, consequently promoting an increase in oil synthesis [59]. In summary, the utilization of mycorrhiza enhances nutrient availability and absorption, providing the necessary energy for oil synthesis. Additionally, the availability of phosphorus facilitates the production of high-energy ATP molecules, further supporting oil synthesis in plants [60]. Additionally, intercropping groundnut with corn decreased the palmitic acid content in peanut oil compared to sole cultivation, thereby enhancing the quality of peanut oil [57]. Research indicates that elevated phosphorus fertilizer consumption led to an increase in the saturated palmitic fatty acid content in peanut oil, resulting in reduced oil quality [57] and an increase in saturated fatty acids in corn [61]. Polyphenol compounds, naturally occurring in plants, possess antioxidant potential and are capable of neutralizing free radicals [62]. The effectiveness of antioxidant activity relies on the arrangement and the number of hydroxyl groups in the phenolic compounds, which can mitigate oxidation by donating hydrogen atoms to radicals [63]. At the Mayamey location, the levels of CAT, POX, and SOD enzymes were found to be 7.50%, 6.25%, and 16.39% higher, respectively, compared to the Shahrood location. Notably, in perennial ryegrass plants, the activity of antioxidant enzymes has been observed to increase through the inoculation with mycorrhiza, indicating a positive correlation between mycorrhizal inoculation and enhanced antioxidant enzyme activity [64]. Polyphenol oxidases, which are copper-containing antioxidant enzymes abundant in plants, utilize oxygen molecules to oxidize orthodiphenolic compounds like caffeic acid and catechol, converting them into quinone [65]. Increased levels of phenolic compounds are associated with elevated antioxidant activity, and tannins specifically exhibit a positive correlation with antioxidant activities [66]. Additionally, the augmentation of catalytic enzyme activity through inoculation with symbiotic fungi has been documented in various plants, including rice [67], roses [68], and citrus [69], aligning with our research findings. Quinoa seeds, in particular, are known to be abundant in bioactive compounds with potent antioxidant properties, making them a valuable source of natural antioxidants [31]. These findings align with the results obtained from our experiment. In the Shahrood region, the highest phenolic component content was observed with mycorrhiza inoculation alone treatment. Conversely, in the Mayamey region, the application of mycorrhiza inoculation combined with phosphorus (50 kg ha -1 ) resulted in the highest observed phenolic component content (Figure 3). Quinoa seeds boast a wealth of protein, lipids, fiber, vitamins, and minerals. Beyond its optimal blend of essential amino acids, quinoa harbors various phytochemicals, such as saponins, phytosterols, phytoecdysteroids, phenolic compounds, polysaccharides, as well as bioactive proteins and peptides. Recent studies showcasing the positive impacts of these compounds on metabolic, cardiovascular, and gastrointestinal health have propelled quinoa into the spotlight as a recognized functional food and nutraceutical [70]. Supplementing phosphorus in conjunction with AMF can modify the availability of nutrients, influencing the metabolic environment of plants and improving their ability to assimilate CO 2 from the atmosphere. For instance, in a study by Kaling et al. [71], transcriptome and metabolome analyses of poplar plants inoculated with mycorrhizae and herbivores revealed that plants exhibit a specific accumulation of specialized protective compounds, such as protease inhibitors and aldoxime, at the cost of plant constitutive phenol-based compounds, as a defense mechanism against herbivores. AMF form a symbiotic relationship with the majority of plants and are typically found on the surface of plant roots and in the vicinity of root cells' epidermis. The hyphae of these fungi have the unique ability to penetrate soil micropores, facilitated by their thinner structure compared to plant root hairs. This enables them to transport minerals, including phosphates and nitrates, from the soil to their host plant in exchange for carbohydrates, as corroborated by the findings of our study and the research conducted by van Der Heijden et al. [72]. While prior studies have noted AMF associations with quinoa, the observed colonization rates have been minimal [73]. In the study by Vestberg et al. [73], a 19% colonization rate was observed, the highest recorded under field conditions before our observation. In greenhouse studies conducted by Kellog et al. [74] on 10 different quinoa genotypes, the reported colonization of quinoa roots by AMF ranged from 0 to 3%. However, Benaffari et al. [18] reported a higher colonization rate of 50%. These findings indicate that the intensity and frequency of AMF colonization in quinoa roots are significantly reduced in a monoculture system (p< 0.05). Conversely, treatments involving intercropping with maize (75% maize: 25% quinoa) and the addition of AMF (AMF+p50 kg ha -1 ) exhibited the highest frequency of mycorrhizal associations. In this study, as the levels of soluble phosphorus in the soil rose, there was a corresponding decrease in the colonization of roots by AMF. These fungi possess specific biochemical and physiological characteristics that enhance the availability of phosphorus to roots. Through the release of protons, these fungi acidify the rhizosphere, thereby increasing the solubility and transfer of phosphorus, particularly in alkaline soils, as observed in the soil of the studied areas. In acidic soils where phosphorus is predominantly bound with iron or aluminum, mycorrhizal fungi's neutralization of chelating agents can enhance the root bioavailability of soil phosphorus [72]. Materials and Methods Within this investigation, a factorial experiment was conducted employing a Randomized Complete Block Design (RCBD) featuring three replications. The investigation was carried out in two distinct locations in Iran, Shahrood and Mayamey, both situated within the Semnan province. Detailed data regarding the geographic and climatic features of the research locations, along with relevant soil physical and chemical attributes, are outlined in Tables 1 and 2. The seeds utilized in this study, including the maize variant 704 and quinoa variety Titicaca, were sourced from the Ministry of Agriculture Jihad in Tehran, Iran. The gathering of plant materials adhered to the supervision and permissions granted by the Ministry of Agriculture Jihad of Tehran, Iran, and conformed to both national and local guidelines. All authors of the study fully adhered to the stipulated local and national guidelines. The cultivation plan consisted of various experimental setups, including the exclusive cultivation of 100% maize, 100% quinoa, and proportional mixtures of both plants (25%, 50%, and 75%). This resulted in five distinct cultivation systems: 100% maize, 100% quinoa, 25% maize: 75% quinoa, 50% maize: 50% quinoa, and 75% maize: 25% quinoa. Each plot measured 4 meters in width and 6 meters in length. The crop rows were oriented in a north-to-south direction. The row spacing and plant spacing were set at 50 cm and 10 cm, respectively. Additionally, there was a distance of 30 cm between the maize and quinoa rows. On May 24, 2022, both quinoa and maize were planted in both locations. Regular irrigation was conducted every six days to maintain adequate moisture levels for the plants. For the standard cultivation of 100% maize and 100% quinoa, four rows of each crop were grown. In the "50% quinoa: 50% maize" system, two rows of quinoa and two rows of maize were cultivated. Similarly, in the "75% maize: 25% quinoa" and "25% maize: 75% quinoa" systems, one row of quinoa was incorporated with three rows of maize and three rows of quinoa with one row of maize, respectively, maintaining the four-row arrangement. Phosphorus fertilizer, in the form of triple superphosphate (Ca (H 2 PO 4 ) 2 H 2 O), was applied at three levels: control (P0), 50 kg ha -1 (P1), and 100 kg ha -1 (P2). The fertilizer was placed at a depth of 5 cm in the soil during planting. Mycorrhiza was utilized in two forms: M1, involving the application of mycorrhiza as a seed coating, and M2, with no mycorrhiza utilization. Minimize any potential marginal effects, samples were collected from the central portion of each plot. The mycorrhizal fungi used in the study were obtained from Green Biotechnology Company located in Karaj, Iran. To enhance the adhesion of the mycorrhizal to the seeds, Arabic gum was employed. The coated seeds were then placed in a polythene bag and vigorously shaken for 30 seconds to ensure a uniform coating on the surface of all the seeds. Subsequently, 50 grams of inoculum per kilogram of seeds was used, and the mixture was thoroughly combined to ensure proper distribution of the fungus. The treated seeds were then dried in the shade for 10 minutes. Following this, the planting operation was swiftly carried out [75]. Measurement of Total Flavonoids Content The aluminum chloride colorimetric method was employed to determine the total flavonoids (TF) content [76]. To begin, 0.05 g of the sample was sonicated in 1.6 mL of 50% methanol for one hour, then placed in a thermostatic water bath oscillator at 65 ° C and 210 rpm for 30 minutes. Following this, the mixture was centrifuged at 10,000 rpm for 10 minutes, and the resulting supernatant was left to incubate at 4 ° C for 24 hours. After combining 2 mL of the extraction solution with 1 mL of 5% NaNO 2 , the mixture was agitated for 6 minutes at 37 ° C. Next, 1 mL of 10% aluminum chloride was introduced into the tube and mixed, followed by the addition of 5 mL of 1 M NaOH with subsequent shaking. The sample's absorbance was measured at 510 nm, with 50% methanol employed as a blank. To ascertain the TF concentration, a calibration curve was formulated using catechol from Sigma Chemicals as the standard. Total Phenolic Compound Content Assay Using the Folin-Ciocalteu colorimetric method [77], the total phenolic compound content (TP) was assessed. This method relies on the reduction of a phosphomolybdic/phosphotungstic acid complex by phenolic compounds in the presence of the Folin-Ciocalteu reagent. To quantify the TP concentration, a calibration curve was formulated utilizing gallic acid from Sigma Chemicals as the reference. The TP value was denoted as milligrams of gallic acid equivalents per gram. Total Saponin Content Assay To determine the total saponin (TS) content, a colorimetric method was implemented [78]. Initially, 0.1 g of leaf extract was subjected to extraction in 4 mL of 75% ethanol using an ultrasonic device for 30 minutes at 30 ° C. After centrifugation at 8000 rpm for 5 minutes, the supernatant was transferred to a tube. The addition of anhydrous methanol adjusted the volume to 5 mL. 0.1 mL of the sample solution was combined with 0.1 mL of the vanillin reagent and 0.4 mL of perchloric acid. After subjecting the mixture to a colorimetric reaction in a water bath at 70 ° C for 15 minutes with vigorous stirring, rapid cooling on ice ensued. Following this, 2.5 mL of glacial acetic acid was introduced, and the blend was chilled in an ice-water bath for 10 minutes. The absorbance of the resulting sample was assessed at 550 nm. The total saponin concentration was ascertained via a calibration curve (y = 0.0534x + 0.0003, R 2 = 0.999) established using oleanolic acid as the blank. TS was quantified in milligrams of oleanolic acid equivalents. Antioxidant Enzymes Catalase Activity A modified technique was utilized for the extraction of antioxidant enzymes [79]. Initially, 0.2 g of plant tissue underwent homogenization with the extraction buffer, including 1600 μl of potassium phosphate buffer (pH = 6.8), 20 μl of 0.1 M EDTA, and 380 μl of distilled water. The ensuing mixture then underwent centrifugation at 4000 rpm for 25 minutes at 4 ° C, and the supernatant phase was utilized for assessing enzyme activity. The spectrophotometric method proposed by Abi [80] was employed to determine the catalase enzyme (CAT) activity. Upon completion of the enzyme extract preparation, a kinetic assay to evaluate catalase activity was initiated. This entailed blending 2.5 ml of potassium phosphate buffer (pH = 7) and 0.3 ml of 3% hydrogen peroxide in an icy bath. Promptly thereafter, 0.2 ml of the enzyme extract was introduced, and the alteration in absorbance at 240 nm was registered via a spectrophotometer. The catalase activity was denoted as mmol of hydrogen peroxide decomposed per minute per milligram of fresh leaf weight (U mg^ -1 fresh leaf weight). Peroxidase Activity For the assessment of peroxidase enzyme (POX) activity, the reaction blend comprised the subsequent concentrations: 1400 μl of 100 mM potassium phosphate buffer (pH = 7), 100 μl of 70 mM soluble potassium phosphate H 2 O 2 (pH = 7), and 750 μl of distilled water. The enzyme extract was blended with this mixture for the assessment of peroxidase activity at 470 nm. The peroxidase activity was computed as mmol of tetraguaiacol generated per minute per milligram of fresh leaf weight, utilizing the approach delineated by Chance and Maehly [81]. Superoxide Dismutase Activity To evaluate the activity of superoxide dismutase (SOD), its capacity to impede the photochemical reduction of NBT (nitro tetrazolium blue chloride) at 560 nm was assessed, following the method pioneered by Beauchamp and Fridovich [82]. A 50 mM phosphate buffer solution at pH 7.5 was formulated. Preparing a 50 mM phosphate buffer solution at pH 7.5, specific volumes of compounds were then added to the buffer to form the reaction mixture. Measurement of Mycorrhizal Colonization Assessing the AMF colonization involved visually inspecting stained roots. Post-cleaning, roots were sectioned into 2.5 cm segments, forming three subsamples, each containing around 20 segments (equivalent to 0.25 g dry weight). In preparation for staining, the roots underwent treatment with hot 10% KOH to eliminate impurities. Subsequently, trypan blue was employed for root staining [83]. The stained roots were arranged on a grid plate and examined at 20 points, with each point scrutinized for the presence of arbuscular vesicles or hyphae [84]. The reported percentage for each structure represents the average proportion of points where these features were observed. Statistical Analysis The examination of this study involved a combined analysis, employing a factorial experimental design. Variance analysis and means comparison were assessed using SAS software (Version 9.4) with a significance level of 5%. MS Excel was utilized for the creation of graphical representations. Declarations Acknowledgments The financial support by Shahrood University of Technology is acknowledged. Author contributions H.A and H.M: Conceptualization, Data curation, Formal analysis, Project administration, Supervision, Validation; Visualization, Writing-Original Draft, Writing -Review & Editing. T.R: Investigation and Methodology, Formal analysis. M.GH and KH.K: Data curation, Validation, Formal analysis and Review & Editing. Ethics approval and consent to participate This article does not involve any studies conducted on human participants or animals by any of the authors. Commercial, or advanced breeding genotypes were used in the study, and no wild material was used. Data availability The data generated or analyzed in this study are included in this article. 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A plant-fungus bioassay supports the classification of quinoa ( Chenopodium quinoa Willd.) as inconsistently mycorrhizal. Microbial ecology. 82 , 135-144.‏ https://doi.org/10.1007/s00248-021-01710-1 (2021). El-Sawah, A. M., El-Keblawy, A., Ali, D. F. I., Ibrahim, H. M., El-Sheikh, M. A., Sharma, A., ... & Sheteiwy, M. S. Arbuscular mycorrhizal fungi and plant growth-promoting rhizobacteria enhance soil key enzymes, plant growth, seed yield, and qualitative attributes of guar. Agriculture. 11(3) , 194.‏ https://doi.org/10.3390/agriculture11030194 (2021). Roberts, J. L., & Moreau, R. Functional properties of spinach (Spinacia oleracea L.) phytochemicals and bioactives. Food & function. 7(8) , 3337-3353.‏ https://doi.org/10.1039/C6FO00051G (2016). Khanam, U. K. S., Oba, S., Yanase, E., & Murakami, Y. Phenolic acids, flavonoids and total antioxidant capacity of selected leafy vegetables. Journal of Functional Foods. 4(4) , 979-987.‏ https://doi.org/10.1016/j.jff.2012.07.006 (2012). Gil-Ramirez, A., Salas-Veizaga, D. M., Grey, C., Karlsson, E. N., Rodriguez-Meizoso, I., & Linares-Pastén, J. A. Integrated process for sequential extraction of saponins, xylan and cellulose from quinoa stalks ( Chenopodium quinoa Willd.). Industrial Crops and Products. 121 , 54-65.‏ https://doi.org/10.1016/j.indcrop.2018.04.074 (2018). Gong, X., Dang, K., Liu, L., Zhao, G., Lv, S., Tian, L., ... & Feng, B. Intercropping combined with nitrogen input promotes proso millet ( Panicum miliaceum L.) growth and resource use efficiency to increase grain yield on the Loess plateau of China. Agricultural Water Management. 243 , 106434.‏ https://doi.org/10.1016/j.agwat.2020.106434 (2021). Aebi, H. [13] Catalase in vitro. In Methods in enzymology. Academic press.‏ 105 ,121-126. https://doi.org/10.1016/S0076-6879(84)05016-3 (1984). Chance, B. Maehly, A.C. Assay of Catalase and Peroxidase. Methods in Enzymology. 2 , 764-775. http://dx.doi.org/10.1016/S0076-6879(55)02300-8 (1995). Beauchamp, C., Fridovich, I. Superoxide dismutase: improved assays and an assay applicable to acrylamide gels. Analytical biochemistry. 44(1) , 276-287.‏ https://doi.org/10.1016/0003-2697(71)90370-8 (1971). Kormanik, P. P. Quantification of vesicular-arbuscular mycorrhizae in plant roots. Methods and principles of mycorrhizal research. 37-46 (1982).‏ Giovannetti, M., & Mosse, B. An evaluation of techniques for measuring vesicular arbuscular mycorrhizal infection in roots. New phytologist. 489-500.‏ http://www.jstor.org/stable/2432123 (1980). Tables Tables 1 to 5 are available in the Supplementary Files section Additional Declarations No competing interests reported. Supplementary Files Tables.docx Cite Share Download PDF Status: Posted Version 1 posted 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-3930581","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":279712753,"identity":"7df3f5bc-ef39-4a26-a2c1-18bbc223fa07","order_by":0,"name":"Tayebe Rostami","email":"","orcid":"","institution":"Shahrood University of Technology","correspondingAuthor":false,"prefix":"","firstName":"Tayebe","middleName":"","lastName":"Rostami","suffix":""},{"id":279712754,"identity":"51b77b4b-545e-413b-a5b6-c786dd61a8b1","order_by":1,"name":"Hamid Abbasdokht","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA20lEQVRIiWNgGAWjYBACAwYeNiDFzCAhwcD4gIHhAElamJkNSNbCJkGUFnP2s8ce/PhjLSc5u/9YNU/NHTl+BuaHj27g0WLZk5du2NuWbiwtc5jtNs+xZ8aSDWzGxjn4HHYgx0yCt+Fw4jyJZKAWtsOJGw7wsEnj1XL+jZnknz8QLcU8/4jRciPHTBpk+GygFmbeNqK0vEs3lgX6RXJGsrHk3L7DxpLNhPxyPvfYwzfAEJO4kfjww5tvh+X42ZsfPsanBQUw8YBIZmKVgwDjD1JUj4JRMApGwYgBAMyUTJNRLyZVAAAAAElFTkSuQmCC","orcid":"","institution":"Shahrood University of Technology","correspondingAuthor":true,"prefix":"","firstName":"Hamid","middleName":"","lastName":"Abbasdokht","suffix":""},{"id":279712755,"identity":"afdcbbd2-4b68-43b2-a61e-a1ea25ce2f20","order_by":2,"name":"Hassan Makarian","email":"","orcid":"","institution":"Shahrood University of Technology","correspondingAuthor":false,"prefix":"","firstName":"Hassan","middleName":"","lastName":"Makarian","suffix":""},{"id":279712756,"identity":"57bb3ec9-3749-49b8-857d-44f18b8c5ea7","order_by":3,"name":"Manoochehr Gholipoor","email":"","orcid":"","institution":"Shahrood University of Technology","correspondingAuthor":false,"prefix":"","firstName":"Manoochehr","middleName":"","lastName":"Gholipoor","suffix":""},{"id":279712757,"identity":"1d43bd0f-86bd-4c05-bf81-7392dcedaf06","order_by":4,"name":"Khalil karimzadehasl","email":"","orcid":"","institution":"Research Institute of Research, Education and Extension of Forests and Rangelands","correspondingAuthor":false,"prefix":"","firstName":"Khalil","middleName":"","lastName":"karimzadehasl","suffix":""}],"badges":[],"createdAt":"2024-02-05 09:59:50","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-3930581/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-3930581/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":52789250,"identity":"431696cf-e29c-4268-aa72-a52828c8e675","added_by":"auto","created_at":"2024-03-15 19:44:55","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":44246,"visible":true,"origin":"","legend":"\u003cp\u003eThe interaction effect of cropping ratio, application of phosphorus, and mycorrhiza on the total flavonoid content in quinoa. Columns sharing identical letters exhibited no significant distinctions.\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-3930581/v1/a3e91a14c087725fa5098a75.png"},{"id":52789219,"identity":"889775dc-60de-4a2c-86e7-ff5a632f1764","added_by":"auto","created_at":"2024-03-15 19:44:45","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":21965,"visible":true,"origin":"","legend":"\u003cp\u003eThe interaction effect of locations, application of phosphorus, and mycorrhiza on total phenolic compound of quinoa leaf. Columns sharing identical letters exhibited no significant distinctions.\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-3930581/v1/c660a3a1834f0e16e96c1a62.png"},{"id":52789221,"identity":"39850f34-fb2c-4299-bfa9-95b212ea743f","added_by":"auto","created_at":"2024-03-15 19:44:46","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":29879,"visible":true,"origin":"","legend":"\u003cp\u003eThe interaction effect of cropping ratio, application of phosphorus, and mycorrhiza on tannin of quinoa. Columns sharing identical letters exhibited no significant distinctions.\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-3930581/v1/f2e94518217fc3a232896c94.png"},{"id":52789244,"identity":"2df247d3-0cf2-4000-b814-33dd35be2362","added_by":"auto","created_at":"2024-03-15 19:44:54","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":33672,"visible":true,"origin":"","legend":"\u003cp\u003eThe interaction effect of cropping ratio, application of phosphorus, and locations on saponin content in quinoa. Columns sharing identical letters exhibited no significant distinctions.\u003c/p\u003e","description":"","filename":"4.png","url":"https://assets-eu.researchsquare.com/files/rs-3930581/v1/f25e64cee6f4c189d2117e45.png"},{"id":52789251,"identity":"a041200d-f0fb-4fb4-b789-a8d3b6f5b700","added_by":"auto","created_at":"2024-03-15 19:44:55","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":31777,"visible":true,"origin":"","legend":"\u003cp\u003eThe interaction effect of cropping ratio, and application of phosphorus on phytic acid content. Columns sharing identical letters exhibited no significant distinctions.\u003c/p\u003e","description":"","filename":"5.png","url":"https://assets-eu.researchsquare.com/files/rs-3930581/v1/9bd206303bbdb5f6cb70779f.png"},{"id":52789236,"identity":"e46835ae-afe4-4cdb-92cc-ffbb9d35d13d","added_by":"auto","created_at":"2024-03-15 19:44:49","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":31110,"visible":true,"origin":"","legend":"\u003cp\u003eThe interaction effect of cropping ratio, phosphorus application and mycorrhiza inoculation on antioxidant activity. Columns sharing identical letters exhibited no significant distinctions\u003c/p\u003e","description":"","filename":"6.png","url":"https://assets-eu.researchsquare.com/files/rs-3930581/v1/8b6a1f0e8326f0fd39274459.png"},{"id":52789235,"identity":"09500bc6-7ec5-43c0-b1f8-68708877ee8c","added_by":"auto","created_at":"2024-03-15 19:44:47","extension":"png","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":32301,"visible":true,"origin":"","legend":"\u003cp\u003eThe interaction effect of cropping ratio, phosphorus application, mycorrhiza inoculation and locations on colonization percentage in quinoa on antioxidant activity. Columns sharing identical letters exhibited no significant distinctions\u003c/p\u003e","description":"","filename":"7.png","url":"https://assets-eu.researchsquare.com/files/rs-3930581/v1/305646d9c00569d43908d697.png"},{"id":59374957,"identity":"9d5ea42d-acaf-4b62-8f25-bf7d79423184","added_by":"auto","created_at":"2024-07-01 03:35:53","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":852490,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-3930581/v1/061d4eba-d201-48a0-a1f8-79514919f757.pdf"},{"id":52789220,"identity":"8ce25969-0cc7-4816-b1e3-8a2fb0c3dcea","added_by":"auto","created_at":"2024-03-15 19:44:45","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":36727,"visible":true,"origin":"","legend":"","description":"","filename":"Tables.docx","url":"https://assets-eu.researchsquare.com/files/rs-3930581/v1/fcc89a1e2a2778cd8ea8b652.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Investigation of Phytochemical Characteristics of Quinoa in an Intercropping Production System with Maize in Response to Mycorrhiza and Phosphorus Fertilizer","fulltext":[{"header":"Introduction","content":"\u003cp\u003eThe United Nations' 2030 Agenda for Sustainable Development emphasizes the importance of establishing sustainable food systems that can provide nutritious diets for a growing global population. This means prioritizing crops that guarantee improved efficiency in using natural resources and contribute to meeting the requirements of a healthy diet. In line with this objective, alternative crops, such as quinoa, have received significant attention [1]. Quinoa (\u003cem\u003eChenopodium quinoa\u003c/em\u003e Willd.) is an annual crop that belongs to the Chenopodiaceae family. It is native to the Andean region in South America and is primarily cultivated in countries such as Peru, Bolivia, Ecuador, Argentina, Colombia, and Chile [2]. According to a report by the Food and Agriculture Organization (FAO), the quinoa cultivation area has expanded from 52,555 hectares in 1961 to 184,585 hectares in 2019. Similarly, quinoa production has risen from 32,435 tons in 1961 to 161,415 tons in 2019 [3]. Quinoa exhibits resilience to agroecological extremes, including variations in soil conditions, rainfall, temperature, and altitude. It can thrive at sea level up to 4500 m above sea level, demonstrating adaptability to frost, drought, and salinity [4]. In contemporary times, quinoa has garnered increased attention due to its exceptional adaptability to diverse environmental conditions, substantial nutritional content, and widespread preference as an ingredient in gluten-free and functional foods. Being gluten-free, it serves as a source of protein, fatty acids, tocopherol, phytosterols, phenolic compounds, and has a low glycemic index [5]. Moreover, quinoa stands out for its abundant natural antioxidants, including ascorbic acid, flavonoids, phenolic compounds, and phytosterols [6, 7].\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eIn today's world, implementing multi-cropping systems has become crucial, particularly in developing countries and among smallholder farmers. This approach allows for the efficient utilization of land and resources, while also reducing the incidence of pests and diseases [8, 9]. Intercropping agricultural systems have demonstrated notable efficacy in boosting production, contingent on specific ecological conditions. Intercropping is particularly preferred in regions with constraints such as limited land availability, labor force, and mechanization, as it enables the optimal utilization of available resources compared to single-cropping systems [10]. Phosphorus is a critical element for the growth and development of plants, playing a vital role in crop production. However, phosphorus deficiency has emerged as a significant constraint on agricultural productivity [11]. Research conducted by Dang et al. [12] on mung bean (\u003cem\u003eVigna radiata\u003c/em\u003e L.) demonstrated that the application of phosphorus fertilizer led to a notable increase in the activities of superoxide dismutase and peroxidase, by 20.9% and 43.7% respectively.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eArbuscular mycorrhizal fungi (AMF) mycelium plays a pivotal role in boosting soil phosphatase and microbial activity, significantly influencing phosphorus availability. In symbiosis with plants, especially crops, AMF fosters root growth and facilitates the conversion of inorganic phosphorus to organic forms [13]. These fungi form symbiotic relationships with various crops, residing both on root surfaces and within the epidermis of root cells.\u0026nbsp;Intercropping soybean and maize, along with AMF inoculation, has shown significant benefits in promoting the growth of both maize and soybean. Furthermore, this approach has led to increased concentrations of available nitrogen and phosphorus in the soil compared to monoculture systems. Additionally, studies have indicated a positive correlation between the availability of phosphorus fertilizer and plant biomass [14]. These findings underscore the importance of phosphorus management strategies in optimizing plant growth and maximizing agricultural productivity. In their study, Amani Machiani et al. [15] found that the application of AMF had a positive impact on the production of secondary metabolites. The researchers specifically investigated the intercropping of soybean and thyme, combined with AMF application. They observed that this intercropping approach led to improvements in both yield and the production of secondary metabolites in thyme [16]. These findings highlight the potential of AMF utilization and intercropping strategies in enhancing the synthesis of valuable secondary metabolites, offering promising avenues for improving crop productivity and quality.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThis study investigated the influence of mycorrhiza inoculation and phosphorus fertilizer application on the phytochemical traits of quinoa within an intercropping system, considering two different climatic conditions. It is noteworthy that the observed colonization of quinoa by mycorrhizal fungi stands as one of the first documented instances of such symbiotic relationships in Iran. The findings of this study contribute to our understanding of how mycorrhiza inoculation and phosphorus fertilizer application can influence the phytochemical composition of quinoa in different environmental conditions.\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003e\u003cstrong\u003eTotal Flavonoids Content\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;In Table 3, the significant influence of various treatments on the phytochemical properties of quinoa is demonstrated. The highest flavonoid content (66.80 mg g\u003csup\u003e-1\u003c/sup\u003e) was observed when cultivating 100% quinoa with mycorrhiza inoculation and applying 50 kg ha\u003csup\u003e-1\u003c/sup\u003e phosphorus at the Mayamey site. On the flip side, the Shahrood location exhibited the lowest flavonoid content (41.5 mg g\u003csup\u003e-1\u003c/sup\u003e) when adopting a planting combination of 25% quinoa + 75% maize, coupled with 100 kg ha\u003csup\u003e-1\u003c/sup\u003e phosphorus application (without mycorrhiza utilization) (Figure 1).\u003c/p\u003e\n\u003cp\u003eTreatments involving mycorrhiza application exhibited superior effectiveness when contrasted with those lacking mycorrhiza inoculation. The diminished flavonoid content in the Shahrood location, particularly in the cultivation blend of 25% quinoa and 75% corn, may be ascribed to the climatic and soil conditions prevalent in this area (Figure 1), and coupled with light shortage due to the shading of corn on quinoa. Furthermore, a positive connection was observed among flavonoids, total phenol, and tannin contents. This implies that the surge in secondary metabolites stems from improved nutrient absorption facilitated by mycorrhiza hyphae (Table 5).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTotal Phenolic Compound (TPC)\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe analysis of variance results reveals a significant impact of intercropping, phosphorus fertilization, and mycorrhiza treatment on the total phenolic compounds content (TPC) of quinoa at two distinct sites (Table 3). The highest TPC value (40.57 mg g\u003csup\u003e-1\u003c/sup\u003e) was detected at the Mayamey location with the combined utilization of 50 kg ha\u003csup\u003e-1\u003c/sup\u003e phosphorus and mycorrhiza, as well as at the Shahrood site with mycorrhiza application and absence of phosphorus. Conversely, the lowest TPC value (29.5 mg g\u003csup\u003e-1\u003c/sup\u003e) was documented at the Shahrood location when applying 100 kg ha\u003csup\u003e-1\u003c/sup\u003e phosphorus (Figure 2).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTannins content\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe influence of varying phosphorus fertilizer levels, geographical locations, and intercropping on tannin levels exhibited a notable significance at a 1% level. Furthermore, all main effects and the interactive effects of cropping ratio, locations, phosphorus fertilizer, and mycorrhizal application were significant at the 1% level (Table 3). Examination of mean comparisons unveiled that the peak tannin content (0.49 mg g \u003csup\u003e-1\u003c/sup\u003e) was identified in the intercropping of 50% quinoa + 50% maize, accompanied by the application of 50 kg ha\u003csup\u003e-1\u003c/sup\u003e phosphorus and mycorrhiza, at the Mayamey location. On the contrary, the least tannin content (0.11 mg g \u003csup\u003e-1\u003c/sup\u003e) was documented under the intercropping of 25% quinoa + 75% maize in the Shahrood locale, in the absence of mycorrhiza utilization (Figure 3). Additionally, a notable positive correlation was observed between the tannin content and the antioxidant activity, alongside antioxidant enzymes (Table 5).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eSaponin content\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe saponin content was significantly influenced (p\u0026lt;0.001) by the three-way interactions involving cropping ratio, locations, and phosphorus fertilizer (Table 3). The results indicated a reduction in saponin content as the ratio of corn increased in the intercropping, evident in both the control group and the application of phosphorus (Figure 4). Additionally, a strong positive correlation at the 1% significance level was found between saponin content and the levels of tannins and flavonoids (Table 5).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003ePhytic acid content\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe phytic acid content in quinoa was significantly impacted by all individual treatments (p\u0026lt;0.001). However, most interactive treatments did not show statistically significant effects on phytic acid content. The treatment with the highest phytic acid content (3.8%) was observed in the intercropping of 50% quinoa and 50% maize with a phosphorus application of 50 kg ha\u003csup\u003e-1\u003c/sup\u003e (Table 3). Conversely, the lowest phytic acid content (2.68%) was found in the intercropping of 25% quinoa + 75% maize in combination with phosphorus application at 100 kg ha\u003csup\u003e-1\u003c/sup\u003e in the Shahrood location (Figure 5).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAntioxidant\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe cropping ratio, phosphorus fertilizer application, mycorrhiza inoculation, and locations had a significant impact on the antioxidant activity in quinoa (Table 3). Upon examining the interactive effects of phosphorus levels with mycorrhiza inoculation, it was found that the highest antioxidant activity (79.85%) was attained with the application of 50 kg ha\u003csup\u003e-1\u003c/sup\u003e of phosphorus, mycorrhiza treatment, and a cropping ratio of 50% quinoa. Conversely, the lowest antioxidant activity (52.05%) was recorded in the intercropping of 25% quinoa ratio, phosphorus application of 100 kg ha\u003csup\u003e-1\u003c/sup\u003e, and the absence of mycorrhiza (Figure 6). Additionally, a significant positive correlation emerged between antioxidant activity and the levels of total phenols, flavonoids, tannins, as well as antioxidant enzymes (CAT, POX, SOD) (Table 5).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAntioxidant Enzymes (CAT, POX, SOD)\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe antioxidant enzyme contents were notably impacted by the cropping ratio, phosphorus application, and mycorrhiza inoculation (P\u0026le;0.05). Elevated levels of antioxidant enzymes were evident with mycorrhizal and phosphorus applications when compared to the control conditions (Table 3). In the Mayamey location, the utilization of mycorrhizal treatment and phosphorus application (50 kg ha\u003csup\u003e-1\u003c/sup\u003e) resulted in higher antioxidant enzyme contents than those observed in the Shahrood location. The highest antioxidant enzyme contents were witnessed with the application of phosphorus fertilizer at 50 kg ha\u003csup\u003e-1\u003c/sup\u003e, while the treatment without phosphorus application and mycorrhiza inoculation exhibited the lowest antioxidant enzyme contents (Table 4).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eColonization Percentage\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe various treatments, encompassing locations, mycorrhiza introduction, phosphorus usage, and intercropping ratio, exhibited a significant influence (p\u0026lt;0.001) on the colonization percentage in quinoa (Table 3). Through mean comparison analysis, it was evident that the highest colonization rate (44.6%) was registered in the intercropping consisting of 25% quinoa and 75% maize, along with the application of 50 kg ha\u003csup\u003e-1\u003c/sup\u003e of phosphorus and the AMF application. Conversely, the lowest colonization rate (5%) was noted in the 100% quinoa cultivation system without phosphorus application (control) in the Mayamey location (Figure 7).\u003c/p\u003e\n\u003cp\u003eA strong and noteworthy correlation was found between the levels of flavonoids and phenols, along with the antioxidant enzyme content (Table 5). In contrast, no significant correlation was identified between phytic acid content and the measured phytochemical properties, encompassing antioxidant levels and related enzymes. The colonization percentage exhibited a significant positive correlation with both flavonoid and phytic acid levels (P\u0026le;0.05). Moreover, no noteworthy correlation was identified between saponin content and the antioxidant enzymes content.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eIn this research, plants subjected to mycorrhizal fungi exhibited an expanded root system, enhancing the availability of water and essential nutrients, notably nitrogen, phosphorus, and potassium, crucial for plant growth and development [17]. The mycorrhizal hyphae played a role in augmenting magnesium absorption, consequently elevating the levels of total chlorophyll and flavonoids in both quinoa and maize plants [18]. The synthesis of total chlorophyll is contingent upon the adequate absorption of minerals, particularly nitrogen and magnesium [19]. Phosphorus plays a crucial physiological role in the biosynthesis pathway of biochemical compositions, as it exhibits a synergistic effect with nitrogen. Our findings indicate that the utilization of AMF resulted in an augmentation of secondary metabolite levels in quinoa compared to the control treatment. This increase may be attributed to the enhanced biochemical compositions [18]. Secondary metabolites, such as flavonoids belonging to phenylpropanoids, play a crucial role in maintaining the structural and functional integrity of cells, thereby mitigating oxidative damage caused by ROS. This protective mechanism aids plants in coping with drought stress [20, 21]. AMF has been observed to significantly enhance anthocyanin and total flavonoid concentrations in \u003cem\u003eMedicago truncatula\u003c/em\u003e leaves [22]. It\u0026apos;s noteworthy that these mycorrhizal effects can, in part, be attributed to the improved phosphorus (P) content [23].\u003c/p\u003e\n\u003cp\u003eXia et al. [21] demonstrated that mycorrhizae lead to the accumulation of root metabolites. However, the exact roles of these secondary metabolites induced by AM symbiosis in enhancing plant drought tolerance remain largely unknown. One of the well-known benefits of AM symbiosis to the host plant is the significant improvement in water and nutrient availability facilitated by AM. This, in turn, can explain the changes observed in carbohydrates and certain components of secondary metabolism, which are referred to as general mycorrhizal effects [24]. While the effects of mycorrhiza on plants are frequently considered to be highly specific to host plants and/or fungal species, there are common characteristics in the alterations of mycorrhiza-associated metabolites in plant roots [25]. In line with our findings, previous studies have reported that mycorrhizal associations often lead to increased levels of bioactive phenolic and lignin metabolites compared to plants not inoculated with AM [26, 27]. Furthermore, mycorrhizal symbiosis can induce plant-specific metabolic changes, particularly in stress conditions, which cannot be fully explained solely by nutrient effects [22].\u0026nbsp;Experimental data and meta-analysis have unveiled highly specific effects of mycorrhizal fungi, plant species, and/or their interactions on plant metabolism [17].\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe growth, development, and metabolic activity of plants are contingent upon their photosynthetic capacity and ability to utilize soil nutrients [28]. By introducing phosphorus supplementation in conjunction with AMF, the levels of available nutrients can be modified, thereby leading to alterations in the metabolic environment of plants. This, in turn, facilitates the enhanced absorption and assimilation of CO\u003csub\u003e2\u003c/sub\u003e from the atmosphere. The presence of mycorrhizal fungi resulted in enhanced biochemical properties, including increased antioxidant enzymes activity, flavonoids content, and total phenol, achieved by reducing peroxidase activity [29]. The hyphae of arbuscular mycorrhizal fungi possess the capability to augment water and nutrient uptake by plants, thereby establishing a symbiotic relationship between plants and AMF [30]. Quinoa seeds are known for their abundance of bioactive compounds, which display potent antioxidant activity and contain natural antioxidant compounds [31]. The total phenolic compounds present in quinoa contribute to its antioxidant activity [32]. According to Nsimba et al. [33], the total phenol content of quinoa extract grown in Japan was measured to be 148.0 mg g\u003csup\u003e-1\u003c/sup\u003e equivalent of tannic acid, which was higher compared to the total phenol content (94.3 mg g\u003csup\u003e-1\u003c/sup\u003e) of quinoa grown in Bolivia. The disparities in phenolic component content between these two quinoa ecotypes can be attributed to variations in environmental factors and agricultural conditions [32]. Similar findings were observed in our experiment. Our results indicated that in the Shahrood region, the highest phenolic component content was observed in the treatment involving mycorrhiza inoculation alone. However, in the Mayamey region, the application of mycorrhiza and 50 kg ha\u003csup\u003e-1\u003c/sup\u003e of phosphorous resulted in the highest phenolic component content (Figure 3).\u003c/p\u003e\n\u003cp\u003eThe total phenolic component content in quinoa seeds varied between 16.8 and 59.7 mg/100 g, whereas the content of soluble phenolic compounds in the samples ranged from 7 to 61 mg/100 g [31].While saponins present in quinoa have been identified as anti-quality factors, isolated saponins also possess intriguing biological properties [34]. Our findings revealed that the saponins content decreased as the proportion of maize in mixed crops increased (Figure 3). The application of 100 kg ha\u003csup\u003e-1\u003c/sup\u003e phosphorus significantly reduced the saponin levels compared to the control and 50 kg ha\u003csup\u003e-1\u003c/sup\u003e phosphorus in the 75% maize proportion. It has been reported that saponins may exhibit nutritional or pharmacological value and their levels can vary under different conditions [35].Tannins, belonging to the group of polyphenols, can form complexes with other macromolecules, leading to various biological effects, both beneficial and undesirable. Polyphenols are natural substances that are widespread and commonly distributed in the plant kingdom [36]. Phenolic compounds, flavonoids, and saponins, which are products of secondary metabolism, exhibit crucial antioxidant activities that promote cardiovascular and cerebrovascular health. Additionally, they possess antiallergenic, antibacterial, anticarcinogenic, and anti-inflammatory effects [37].\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eIntercropping systems in agriculture have demonstrated significant effectiveness in enhancing production, contingent upon specific ecological conditions [38]. In a study conducted by Koca [39] on the forage yield and growth parameters of the maize-quinoa intercropping system, it was observed that the amount of maize dry matter increased in nearly all mixtures. Li and Cai [28] observed that enhancing the presence of AMF led to significant improvements in various maize traits, including biomass, chlorophyll content, plant height, and leaf area. AMF mycelium plays a crucial role in enhancing phosphatase activity and microbial activity in the soil. These fungi have a significant impact on the availability of phosphorus in the soil. Through their symbiotic relationship with plants, particularly crops, AMF promotes root growth and facilitates the conversion of inorganic phosphorus into organic forms [13]. AMF can be found in symbiosis with a wide range of crops, residing on the surface of roots and within the epidermis of root cells. The hyphae of mycorrhizal fungi are finer than root hairs, allowing them to penetrate soil micropores and transport certain minerals such as nitrates and phosphates to their host plant in exchange for carbohydrates [40]. This aligns with the findings of the current study. Consequently, plants associated with mycorrhizae generally exhibit greater drought tolerance compared to those without mycorrhizal associations [41]. Phenolic compounds, as secondary metabolites, play a role in detoxification, aiding in the neutralization of active oxygen species and free radicals before they can harm plant cells. The antioxidant potential of a plant is contingent upon the concentration of phenolic and tannin compounds. This study demonstrated an increase in the amount of phenolic and tannin compounds when AMF were utilized. Under stress conditions, mycorrhizal plants respond to oxidative stress by enhancing the synthesis of phenolic compounds. Previous research has also reported an increase in phenolic and tannin compounds facilitated by AMF in various plants, including violets [42], grapes [43], and \u003cem\u003eCommiphora leptophloeos\u003c/em\u003e [44].\u003c/p\u003e\n\u003cp\u003eIn our study, variations in the saponin content of quinoa were observed. As per the results, the saponin content decreased with an increase in the proportion of maize in mixed crops (Figure 3). The application of 100 kg ha\u003csup\u003e-1\u003c/sup\u003e phosphorus significantly reduced the saponin content compared to the control and 50 kg ha\u003csup\u003e-1\u003c/sup\u003e phosphorus in the 75% maize proportion. Additionally, when quinoa samples underwent different pearling processes, the saponin levels decreased [45]. These findings highlight the influence of cropping proportion, phosphorus application, and processing techniques on the saponin content of quinoa. Moreover, the quantities and composition of saponins in the quinoa plant were not consistently stable due to the continual removal of seed pericarps and bran in the harvesting process to render it edible, for instance, through pressure-cooking and toasting [45]. The total saponin content in quinoa grown ranged from 3.81 to 27.1 mg gr\u003csup\u003e-1\u003c/sup\u003e in Washington State [46, 47], aligning with our findings. An increase in the nitrogen (N) uptake rate per root length has been shown to enhance saponin accumulation in roots, indicating a significant positive relationship between N uptake rate per root length and saponin content in taproots. Sufficient nitrogen fertilizer has the potential to enhance both root structure and nutrient uptake efficiency, subsequently promoting saponin synthesis [48]. Given the crucial role of nitrogen in saponin synthesis, the reduction in saponin content observed in high concentrations of phosphorus (100 kg ha\u003csup\u003e-1\u003c/sup\u003e) in this study may be attributed to a decrease in nitrogen absorption and, consequently, a decrease in saponin contents.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eAndrino et al. [49] demonstrated the presence of mono-, di-, and tricarboxylic low-molecular-weight organic acids in compartments containing orthophosphate (OP) or goethite-bound-orthophosphate (GOE-PA) and phytic acid (PA or GOE-PA), indicating the occurrence of reductive dissolution and ligand exchange/dissolution reactions. Additionally, hyphae grown in goethite loaded with OP and PA exhibited an elevated content of unsaturated lipids, suggesting increased membrane fluidity to maintain optimal hyphal functionality and facilitate phosphorus incorporation. In the presence of plants, AMF demonstrates extensive practical diversity and contributes to growth-promoting functions by producing various metabolites. These functions include the mineralization of phytate, the production of siderophores, the dissolution of phosphorus, and the production of low-molecular-weight organic acids [50]. Additionally, the mycelium of rhizobium plays a role in mobilizing phosphorus from acid phytic, as highlighted by Selvakumar et al. [51]. \u0026nbsp;AMF contribute to phytate mineralization, leading to the transfer of phosphorus [52]. These studies collectively emphasize the significant contributions of AMF in promoting plant growth by facilitating the mineralization of phytate and enhancing phosphorus availability through various mechanisms. While previous studies have predominantly focused on the mechanisms of interaction between plants and beneficial microbes, such as bacteria and AMF, with an emphasis on model and crop plants [53, 25], the associated transcriptomic and metabolomic changes induced by AMF to alleviate effects of adverse environmental conditions in non-model plant species have not been fully elucidated. The photosynthetic capacity and ability to utilize soil nutrients are crucial factors that influence the growth, development, metabolic activities, and fatty acid composition of plants [28]. Zamani et al. [54] demonstrated that the amounts of phenolic compounds and fatty acid profiles in the intercropping of \u003cem\u003eLallemantia iberica\u003c/em\u003e and \u003cem\u003eCicer arietinum\u003c/em\u003e L. increased with the application of AMF.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eIn the 50% intercropping pattern of quinoa with the application of AMF, the levels of phytic acid in quinoa increased. This can be attributed to the improved accessibility of nutrients facilitated by AMF, which in turn affects plant growth and photosynthesis. These changes in nutrient availability and metabolic processes can influence the production of fatty acid precursor compounds and the activity of enzymes like fatty acid synthase and acetyl-CoA carboxylase [55]. Furthermore, other studies have highlighted the positive effects of intercropping systems on plant nutrient availability, oil productivity, and oil quality. For instance, Rezaei Chiyaneh et al. [56] demonstrated that a cropping ratio of 50:50 in an intercrop of black cumin with fenugreek, using biofertilizer, resulted in the highest oil concentration in black cumin. This increase in oil concentration can be attributed to the enhanced nutrient uptake facilitated by the intercropping system. These findings emphasize the potential benefits of incorporating AMF and intercropping practices with biofertilizers in enhancing nutrient availability, oil production, and oil quality in various plant species. Rezapour et al. [57] demonstrated that the joint application of phosphorus and AMF resulted in an enhanced synthesis of fatty acids in groundnut. Additionally, the application of phosphorus fertilizer in a groundnut-corn intercropping system with a ratio of 1:2 improved groundnut oil content [57]. In a parallel study, it was found that the combined use of phosphorus chemical fertilizer and biological fertilizer increased safflower oil yield. These findings highlight the potential of using phosphorus fertilizers in conjunction with AMF and intercropping systems to enhance fatty acid synthesis and improve the oil content and yield of various oilseed crops like groundnut and safflower [58].\u003c/p\u003e\n\u003cp\u003eOil synthesis, compared to other compounds studied in this research, requires a greater amount of energy. As a result, the utilization of mycorrhiza plays a significant role in enhancing the availability and absorption of nutrients by the plant root, providing the necessary energy for oil synthesis. This increased nutrient uptake facilitated by mycorrhiza ultimately leads to an augmentation in the production of oil. Furthermore, the energy required by plant processes is supplied in the form of high-energy ATP molecules. Phosphorus plays a crucial role in the formation of these energy-rich molecules. Therefore, the availability of phosphorus in the plant system contributes to the production of more high-energy ATP molecules, consequently promoting an increase in oil synthesis [59]. In summary, the utilization of mycorrhiza enhances nutrient availability and absorption, providing the necessary energy for oil synthesis. Additionally, the availability of phosphorus facilitates the production of high-energy ATP molecules, further supporting oil synthesis in plants [60]. Additionally, intercropping groundnut with corn decreased the palmitic acid content in peanut oil compared to sole cultivation, thereby enhancing the quality of peanut oil [57]. Research indicates that elevated phosphorus fertilizer consumption led to an increase in the saturated palmitic fatty acid content in peanut oil, resulting in reduced oil quality [57] and an increase in saturated fatty acids in corn [61]. Polyphenol compounds, naturally occurring in plants, possess antioxidant potential and are capable of neutralizing free radicals [62]. The effectiveness of antioxidant activity relies on the arrangement and the number of\u003cspan dir=\"RTL\"\u003e\u0026nbsp;\u003c/span\u003ehydroxyl groups in the phenolic compounds, which can mitigate oxidation by donating hydrogen atoms to radicals [63]. At the Mayamey location, the levels of CAT, POX, and SOD enzymes were found to be 7.50%, 6.25%, and 16.39% higher, respectively, compared to the Shahrood location. Notably, in perennial ryegrass plants, the activity of antioxidant enzymes has been observed to increase through the inoculation with mycorrhiza, indicating a positive correlation between mycorrhizal inoculation and enhanced antioxidant enzyme activity [64]. Polyphenol oxidases, which are copper-containing antioxidant enzymes abundant in plants, utilize oxygen molecules to oxidize orthodiphenolic compounds like caffeic acid and catechol, converting them into quinone [65]. Increased levels of phenolic compounds are associated with elevated antioxidant activity, and tannins specifically exhibit a positive correlation with antioxidant activities [66]. Additionally, the augmentation of catalytic enzyme activity through inoculation with symbiotic fungi has been documented in various plants, including rice [67], roses [68], and citrus [69], aligning with our research findings. Quinoa seeds, in particular, are known to be abundant in bioactive compounds with potent antioxidant properties, making them a valuable source of natural antioxidants [31]. These findings align with the results obtained from our experiment. \u0026nbsp;In the Shahrood region, the highest phenolic component content was observed with mycorrhiza inoculation alone treatment. Conversely, in the Mayamey region, the application of mycorrhiza inoculation combined with phosphorus (50 kg ha\u003csup\u003e-1\u003c/sup\u003e) resulted in the highest observed phenolic component content (Figure 3). Quinoa seeds boast a wealth of protein, lipids, fiber, vitamins, and minerals. Beyond its optimal blend of essential amino acids, quinoa harbors various phytochemicals, such as saponins, phytosterols, phytoecdysteroids, phenolic compounds, polysaccharides, as well as bioactive proteins and peptides. Recent studies showcasing the positive impacts of these compounds on metabolic, cardiovascular, and gastrointestinal health have propelled quinoa into the spotlight as a recognized functional food and nutraceutical [70]. Supplementing phosphorus in conjunction with AMF can modify the availability of nutrients, influencing the metabolic environment of plants and improving their ability to assimilate CO\u003csub\u003e2\u003c/sub\u003e from the atmosphere. For instance, in a study by Kaling et al. [71], transcriptome and metabolome analyses of poplar plants inoculated with mycorrhizae and herbivores revealed that plants exhibit a specific accumulation of specialized protective compounds, such as protease inhibitors and aldoxime, at the cost of plant constitutive phenol-based compounds, as a defense mechanism against herbivores. AMF form a symbiotic relationship with the majority of plants and are typically found on the surface of plant roots and in the vicinity of root cells\u0026apos; epidermis. The hyphae of these fungi have the unique ability to penetrate soil micropores, facilitated by their thinner structure compared to plant root hairs. This enables them to transport minerals, including phosphates and nitrates, from the soil to their host plant in exchange for carbohydrates, as corroborated by the findings of our study and the research conducted by van Der Heijden et al. [72].\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eWhile prior studies have noted AMF associations with quinoa, the observed colonization rates have been minimal [73]. In the study by Vestberg et al. [73], a 19% colonization rate was observed, the highest recorded under field conditions before our observation. In greenhouse studies conducted by Kellog et al. [74] on 10 different quinoa genotypes, the reported colonization of quinoa roots by AMF ranged from 0 to 3%. However, Benaffari et al. [18] reported a higher colonization rate of 50%. These findings indicate that the intensity and frequency of AMF colonization in quinoa roots are significantly reduced in a monoculture system (p\u0026lt; 0.05). Conversely, treatments involving intercropping with maize (75% maize: 25% quinoa) and the addition of AMF (AMF+p50 kg ha\u003csup\u003e-1\u003c/sup\u003e) exhibited the highest frequency of mycorrhizal associations. In this study, as the levels of soluble phosphorus in the soil rose, there was a corresponding decrease in the colonization of roots by AMF. These fungi possess specific biochemical and physiological characteristics that enhance the availability of phosphorus to roots. Through the release of protons, these fungi acidify the rhizosphere, thereby increasing the solubility and transfer of phosphorus, particularly in alkaline soils, as observed in the soil of the studied areas. In acidic soils where phosphorus is predominantly bound with iron or aluminum, mycorrhizal fungi\u0026apos;s neutralization of chelating agents can enhance the root bioavailability of soil phosphorus [72].\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cp\u003eWithin this investigation, a factorial experiment was conducted employing a Randomized Complete Block Design (RCBD) featuring three replications. The investigation was carried out in two distinct locations in Iran, Shahrood and Mayamey, both situated within the Semnan province. Detailed data regarding the geographic and climatic features of the research locations, along with relevant soil physical and chemical attributes, are outlined in Tables 1 and 2.\u003c/p\u003e\n\u003cp\u003eThe seeds utilized in this study, including the maize variant 704 and quinoa variety Titicaca, were sourced from the Ministry of Agriculture Jihad in Tehran, Iran. The gathering of plant materials adhered to the supervision and permissions granted by the Ministry of Agriculture Jihad of Tehran, Iran, and conformed to both national and local guidelines. All authors of the study fully adhered to the stipulated local and national guidelines. \u0026nbsp;The cultivation plan consisted of various experimental setups, including the exclusive cultivation of 100% maize, 100% quinoa, and proportional mixtures of both plants (25%, 50%, and 75%). This resulted in five distinct cultivation systems: 100% maize, 100% quinoa, 25% maize: 75% quinoa, 50% maize: 50% quinoa, and 75% maize: 25% quinoa. Each plot measured 4 meters in width and 6 meters in length. The crop rows were oriented in a north-to-south direction. The row spacing and plant spacing were set at 50 cm and 10 cm, respectively. Additionally, there was a distance of 30 cm between the maize and quinoa rows. On May 24, 2022, both quinoa and maize were planted in both locations. Regular irrigation was conducted every six days to maintain adequate moisture levels for the plants. For the standard cultivation of 100% maize and 100% quinoa, four rows of each crop were grown. In the \"50% quinoa: 50% maize\" system, two rows of quinoa and two rows of maize were cultivated. Similarly, in the \"75% maize: 25% quinoa\" and \"25% maize: 75% quinoa\" systems, one row of quinoa was incorporated with three rows of maize and three rows of quinoa with one row of maize, respectively, maintaining the four-row arrangement. Phosphorus fertilizer, in the form of triple superphosphate (Ca (H\u003csub\u003e2\u003c/sub\u003ePO\u003csub\u003e4\u003c/sub\u003e)\u003csub\u003e2\u0026nbsp;\u003c/sub\u003eH\u003csub\u003e2\u003c/sub\u003eO), was applied at three levels: control (P0), 50 kg ha\u003csup\u003e-1\u003c/sup\u003e (P1), and 100 kg ha\u003csup\u003e-1\u003c/sup\u003e (P2). The fertilizer was placed at a depth of 5 cm in the soil during planting. Mycorrhiza was utilized in two forms: M1, involving the application of mycorrhiza as a seed coating, and M2, with no mycorrhiza utilization. Minimize any potential marginal effects, samples were collected from the central portion of each plot. The mycorrhizal fungi used in the study were obtained from Green Biotechnology Company located in Karaj, Iran. To enhance the adhesion of the mycorrhizal to the seeds, Arabic gum was employed. The coated seeds were then placed in a polythene bag and vigorously shaken for 30 seconds to ensure a uniform coating on the surface of all the seeds. Subsequently, 50 grams of inoculum per kilogram of seeds was used, and the mixture was thoroughly combined to ensure proper distribution of the fungus. The treated seeds were then dried in the shade for 10 minutes. Following this, the planting operation was swiftly carried out [75].\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eMeasurement of Total Flavonoids Content\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe aluminum chloride colorimetric method was employed to determine the total flavonoids (TF) content [76]. To begin, 0.05 g of the sample was sonicated in 1.6 mL of 50% methanol for one hour, then placed in a thermostatic water bath oscillator at 65 \u003csup\u003e°\u003c/sup\u003eC and 210 rpm for 30 minutes. Following this, the mixture was centrifuged at 10,000 rpm for 10 minutes, and the resulting supernatant was left to incubate at 4 \u003csup\u003e°\u003c/sup\u003eC for 24 hours. After combining 2 mL of the extraction solution with 1 mL of 5% NaNO\u003csub\u003e2\u003c/sub\u003e, the mixture was agitated for 6 minutes at 37 \u003csup\u003e°\u003c/sup\u003eC. Next, 1 mL of 10% aluminum chloride was introduced into the tube and mixed, followed by the addition of 5 mL of 1 M NaOH with subsequent shaking. The sample's absorbance was measured at 510 nm, with 50% methanol employed as a blank. To ascertain the TF concentration, a calibration curve was formulated using catechol from Sigma Chemicals as the standard.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTotal Phenolic Compound Content Assay\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eUsing the Folin-Ciocalteu colorimetric method [77], the total phenolic compound content (TP) was assessed. This method relies on the reduction of a phosphomolybdic/phosphotungstic acid complex by phenolic compounds in the presence of the Folin-Ciocalteu reagent. To quantify the TP concentration, a calibration curve was formulated utilizing gallic acid from Sigma Chemicals as the reference. The TP value was denoted as milligrams of gallic acid equivalents per gram.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTotal Saponin Content Assay\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eTo determine the total saponin (TS) content, a colorimetric method was implemented [78]. Initially, 0.1 g of leaf extract was subjected to extraction in 4 mL of 75% ethanol using an ultrasonic device for 30 minutes at 30 \u003csup\u003e°\u003c/sup\u003eC. After centrifugation at 8000 rpm for 5 minutes, the supernatant was transferred to a tube. The addition of anhydrous methanol adjusted the volume to 5 mL. 0.1 mL of the sample solution was combined with 0.1 mL of the vanillin reagent and 0.4 mL of perchloric acid. After subjecting the mixture to a colorimetric reaction in a water bath at 70 \u003csup\u003e°\u003c/sup\u003eC for 15 minutes with vigorous stirring, rapid cooling on ice ensued. Following this, 2.5 mL of glacial acetic acid was introduced, and the blend was chilled in an ice-water bath for 10 minutes. The absorbance of the resulting sample was assessed at 550 nm. The total saponin concentration was ascertained via a calibration curve (y = 0.0534x + 0.0003, R\u003csup\u003e2\u003c/sup\u003e = 0.999) established using oleanolic acid as the blank. TS was quantified in milligrams of oleanolic acid equivalents.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAntioxidant Enzymes\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCatalase Activity\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eA modified technique was utilized for the extraction of antioxidant enzymes [79]. Initially, 0.2 g of plant tissue underwent homogenization with the extraction buffer, including 1600 μl of potassium phosphate buffer (pH = 6.8), 20 μl of 0.1 M EDTA, and 380 μl of distilled water. The ensuing mixture then underwent centrifugation at 4000 rpm for 25 minutes at 4 \u003csup\u003e°\u003c/sup\u003eC, and the supernatant phase was utilized for assessing enzyme activity. The spectrophotometric method proposed by Abi [80] was employed to determine the catalase enzyme (CAT) activity. Upon completion of the enzyme extract preparation, a kinetic assay to evaluate catalase activity was initiated. This entailed blending 2.5 ml of potassium phosphate buffer (pH = 7) and 0.3 ml of 3% hydrogen peroxide in an icy bath. Promptly thereafter, 0.2 ml of the enzyme extract was introduced, and the alteration in absorbance at 240 nm was registered via a spectrophotometer. The catalase activity was denoted as mmol of hydrogen peroxide decomposed per minute per milligram of fresh leaf weight (U mg^\u003csup\u003e-1\u003c/sup\u003e fresh leaf weight).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003ePeroxidase Activity\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eFor the assessment of peroxidase enzyme (POX) activity, the reaction blend comprised the subsequent concentrations: 1400 μl of 100 mM potassium phosphate buffer (pH = 7), 100 μl of 70 mM soluble potassium phosphate H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u0026nbsp;\u003c/sub\u003e(pH = 7), and 750 μl of distilled water. The enzyme extract was blended with this mixture for the assessment of peroxidase activity at 470 nm. The peroxidase activity was computed as mmol of tetraguaiacol generated per minute per milligram of fresh leaf weight, utilizing the approach delineated by Chance and Maehly [81].\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eSuperoxide Dismutase Activity\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eTo evaluate the activity of superoxide dismutase (SOD), its capacity to impede the photochemical reduction of NBT (nitro tetrazolium blue chloride) at 560 nm was assessed, following the method pioneered by Beauchamp and Fridovich [82]. A 50 mM phosphate buffer solution at pH 7.5 was formulated. Preparing a 50 mM phosphate buffer solution at pH 7.5, specific volumes of compounds were then added to the buffer to form the reaction mixture.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eMeasurement of Mycorrhizal Colonization\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAssessing the AMF colonization involved visually inspecting stained roots. Post-cleaning, roots were sectioned into 2.5 cm segments, forming three subsamples, each containing around 20 segments (equivalent to 0.25 g dry weight). In preparation for staining, the roots underwent treatment with hot 10% KOH to eliminate impurities. Subsequently, trypan blue was employed for root staining [83]. The stained roots were arranged on a grid plate and examined at 20 points, with each point scrutinized for the presence of arbuscular vesicles or hyphae [84]. The reported percentage for each structure represents the average proportion of points where these features were observed.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eStatistical Analysis \u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe examination of this study involved a combined analysis, employing a factorial experimental design. \u0026nbsp;Variance analysis and means comparison were assessed using SAS software (Version 9.4) with a significance level of 5%. MS Excel was utilized for the creation of graphical representations.\u0026nbsp;\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgments\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;The financial support by Shahrood University of Technology is acknowledged.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;H.A and H.M: Conceptualization, Data curation, Formal analysis, Project administration, Supervision, Validation; Visualization, Writing-Original Draft, Writing -Review \u0026amp; Editing. T.R: Investigation and Methodology, Formal analysis. M.GH and KH.K: Data curation, Validation, Formal analysis and Review \u0026amp; Editing.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEthics approval and consent to participate\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis article does not involve any studies conducted on human participants or animals by any of the authors. Commercial, or advanced breeding genotypes were used in the study, and no wild material was used.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData availability\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe data generated or analyzed in this study are included in this article. Other materials that support the findings of this study are available from the corresponding author on reasonable request.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent for publication\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting interests\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare no competing interests.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003ePernet, C. A., Ribi Forclaz, A. Revisiting the Food and Agriculture Organization (FAO): international histories of agri-culture, nutrition, and development. The International History Review. \u003cstrong\u003e41(2)\u003c/strong\u003e, 345-350.\u0026rlm; https://doi.org/10.1080/07075332.2018.1460386 (2019).\u003c/li\u003e\n\u003cli\u003eBazile, D., Jacobsen, S. E., Verniau, A. 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New phytologist. 489-500.\u0026rlm; http://www.jstor.org/stable/2432123 (1980). \u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Tables","content":"\u003cp\u003eTables 1 to 5 are available in the Supplementary Files section\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Mixed cropping, Quinoa, Arbuscular mycorrhizal fungi, Phytochemical traits and Phosphorus fertilizer.","lastPublishedDoi":"10.21203/rs.3.rs-3930581/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-3930581/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"Incorporating bio-fertilizers, such as arbuscular mycorrhizal fungi (AMF), into mixed cropping systems has emerged as an innovative and sustainable approach to enhance crop growth and promote sustainable production. The study was conducted in two distinct locations within a mixed cropping system of corn and quinoa. Phosphorus fertilizer was applied at three different levels 50, 100, and 150 kg/ha while also considering the presence or absence of mycorrhiza. Cropping ratios included individual cultivation of quinoa, maize, and intercrop ratios of 50: 50, 75: 25, and 25: 75 (maize: quinoa). The utilization of phosphorus fertilizer improved the symbiotic interaction between plants and arbuscular mycorrhizal fungi, resulting in an increased rate of inoculation by AMF. The 50: 50 intercropping ratio, combined with the application of AMF, resulted in the highest contents of flavonoids, total phenolic compounds, tannins, saponin, antioxidant activity, and phytic acid in quinoa. Our findings demonstrated that AMF exhibit specific roles in optimizing plant strategies for phosphorus uptake and the efficient utilization of phosphorus in the mixed cropping. Recommending the application of AMF in a 50:50 intercropping ratio could be proposed to farmers as a friendly method to attain favorable phytochemical characteristics, particularly when using 50 kg/ha of phosphorus fertilizer.","manuscriptTitle":"Investigation of Phytochemical Characteristics of Quinoa in an Intercropping Production System with Maize in Response to Mycorrhiza and Phosphorus Fertilizer","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-03-15 19:44:19","doi":"10.21203/rs.3.rs-3930581/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"908611a2-38b0-4c6d-9489-7097291ba98a","owner":[],"postedDate":"March 15th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[{"id":29459738,"name":"Biological sciences/Chemical biology"},{"id":29459739,"name":"Biological sciences/Ecology"},{"id":29459740,"name":"Biological sciences/Physiology"}],"tags":[],"updatedAt":"2024-07-01T03:27:46+00:00","versionOfRecord":[],"versionCreatedAt":"2024-03-15 19:44:19","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-3930581","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-3930581","identity":"rs-3930581","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}