Arbuscular mycorrhizal fungus Acaulospora delicata modulates tri-trophic interactions between wheat cultivars and Sitobion avenae under drought stress

Arbuscular mycorrhizal fungus Acaulospora delicata modulates tri-trophic interactions between wheat cultivars and Sitobion avenae under drought stress | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Arbuscular mycorrhizal fungus Acaulospora delicata modulates tri-trophic interactions between wheat cultivars and Sitobion avenae under drought stress Abdul Ghaffar Khoso, Muhammad Awais This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8715853/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 Aphid-microbe-plant interactions are fundamental to understanding plant responses to biotic and abiotic stressors. Aphid is a significant pest distressing wheat crops globally, especially under drought conditions. The interactions between the aphid Sitobion avenae and arbuscular mycorrhizal fungus (AMF) Acaulospora delicata , particularly in wheat cultivars under drought conditions, reveal significant dynamics affecting pest performance and plant health. However, the influence of AMF on aphid performance across different wheat-cultivars under water-deficit stress, remains poorly understood. We investigated the effects of AMF on the performance of S. avenae on two wheat cultivars, Yunhao-618 (drought-resistant) and Xinong-1376 (drought-non-resistant), under varying water conditions. Our results revealed that A. delicata significantly increased root length colonization in Xinong-1376 (67%) and improves aphid developmental duration, longevity, and fecundity under both well-watered and water-deficit conditions. Notably, aphid nymphs exhibited prolonged DDs without AMF-associations, particularly on Xinong-1376 under drought-stress. In contrast, associations with AMF promoted faster growth rates and higher fecundity in Yunhao-618, suggesting that AMF-association can improve water stress effects. Honeydew production was higher in AMF-associated plants under well-watered conditions. Aphid fresh and dry body masses, and water balance traits tended higher on drought-resistant cultivars Yunhao-618 with AMF association under well-water conditions. Additionally, aphids displayed a preference for AMF-associated drought non-resistant Xinong-1376 plants, indicating a strong influence of AMF on host choice dynamics. Our findings reveal that A. delicata modulates plant-aphid dynamics by enhancing plant drought resilience while inadvertently promoting aphid fitness. This underscores the complex role of AMF intri-trophic interactions and their broader implications under climate change scenarios. Entomology Mycorrhiza aphid life-history host choice water-stress AMF-plant-aphid interactions Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 Figure 9 Figure 10 Figure 11 Figure 12 Introduction Climate change is making droughts more frequent, persistent and severe, exacerbating droughts caused by insufficient rainfall or snowfall, leading to water scarcity, soil drought and crop damage (NASA 2023 ; UNICEF 2024 ). These are the main causes of drought stress and the most critical abiotic stresses affecting global agricultural production such as wheat, with crop yields falling by 60% or more in hard-hit areas (Sareen et al. 2023 ; Nyaupane et al. 2024 ). Wheat ( Triticum aestivum ) is a major cereal crop that is important to the world's food supply and nutritional security for several hundred million people around the world (Rakszegi et al. 2023 ; Endalew et al. 2023 ). Wheat is highly susceptible to water scarcity, which can alter its physiological and metabolic processes, leading to growth damage, reduced biomass, and decreased grain quality (Filip et al. 2023 ; Elnajar et al. 2024 ). Wheat crops are also susceptible to various biotic stresses, including insect pests. Aphids, especially wheat aphids, are one of the most destructive pests of wheat, causing direct damage through feeding on the phloem and indirect damage through vector plant viruses such as barley yellow dwarf virus (Ding et al. 2016 ; Leybourne et al. 2024 ) and excretes a sticky substance called honeydew, which is a food source for sooty mold and promotes their growth. Sooty mold is a black fungus that appears on the ears, rather than on other surfaces of wheat leaves and plants (Sabri et al. 2013 ; Talabac 2022 ; Lee 2024 ), significantly reduced their photosynthetic capacity, thereby harming the health of plants. Aphid infestation can significantly reduce wheat productivity, with yield losses ranging from 10% to 70%, depending on the severity of the infection and the health status of the plant (Yahya et al. 2017 ). Arbuscular mycorrhizal fungi (AMF) have established widespread attention for their potential to enhance plant tolerance to abiotic, biotic stresses (Ahammed et al. 2023 ) and forms a binding relationship with plant roots, extending hyphae into the soil and promoting greater water and nutrient absorption (Liu et al. 2018b ). In return, the plants provide the fungi with carbohydrates produced via photosynthesis (Gunjal 2023 ). In addition to their role in nutrient acquisition, AM fungi have been shown to enhance plant drought tolerance by increasing root hydraulic conductivity, enhancing osmotic regulation, and strengthening the synthesis of drought related hormones (Diagne et al. 2020 ; Ye et al. 2023 ). The role of AMF affect plant defense against herbivores by regulating primary and secondary metabolic pathways in plants (Amani et al. 2022 ; Orine et al. 2022 ). AMF, Acaulospora delicata , is a group of fungi that inhabit soil and establish reciprocal relationships with the roots of most terrestrial plants, including wheat, to increase resistance to various abiotic and biotic stresses. For instance, wheat plants colonized by A. delicata are better equipped to cope with drought conditions due to increased nutrient uptake, particularly Phosphorus, and improved water uptake and retention through an extensive hyphal network (da Trindade et al. 2019 ). The colonization may affect the interaction between plants and pests by altering the plants’ nutritional status and defense mechanisms, thereby increasing or decreasing their susceptibility to herbivores such as S. avenae (Koricheva et al. 2009). However, the effects of A. delicata on wheat's interactions with S. avenae under water-stress conditions have not been adequately studied. It is crucial to study the role of AMF, especially A. delicata , in regulating wheat aphid interactions under drought conditions for several reasons. Firstly, it can provide insights into sustainable pest management strategies by utilizing natural plant fungal associations rather than relying solely on chemical insecticides. This sustainable approach is increasingly necessary due to growing concerns about pesticide resistance among pests and the negative environmental impacts associated with chemical use (Zou et al. 2021 ). Secondly, understanding the comprehensive impact of drought resistant cultivars and AMF on aphid performance will provide information for cultivating more stress resistant wheat cultivars. These cultivars may utilize the benefits of AMF combination to enhance drought resistance and resistance to aphid infestation (Madouh et al. 2023). Thirdly, this study will contribute to a broader understanding of how global climate change will affect the dynamics of plant pests, and it is expected that global climate change will increase the frequency and severity of droughts. Investigating the interactions between AMF, wheat, and aphids under conditions of water scarcity will enable better predictions and mitigation strategies for the possible impacts of climate change on agricultural systems (Tang et al. 2022 ; George and Ray 2023 ). Finally, the findings from this study could have broader implications across other cereal crops and pest interactions. Understanding the mechanisms that govern AMF-plant-insect interactions may reveal similarities that can be applied to various species, facilitating enhanced agricultural practices (Pons et al. 2020 ; Wahab et al. 2023 ). The interaction between wheat cultivars, pests, and water stress is complex (Khoso et al. 2025a ). Drought resistant wheat cultivars may enhance their defense against pests such as aphids, possibly due to physiological and morphological adaptations to water stress (Li et al. 2024 ). However, it remains unclear how biotic interactions, particularly with aphids, shift when water stress is involved. The role of associative relationships, such as the association between wheat roots and AMF, under these stressful conditions has been limitedly explored (Rahman et al. 2024 ). It is known that A. delicata enhances plant resistance to abiotic stress by increasing nutrient and water absorption (Carreón-Abud and Gavito 2021 ; Zhang et al. 2022 ), but its role in regulating wheat aphid interactions under drought stress is not fully understood. This study aims to explore the effects of wheat cultivars and A. delicata on the performance of S. avenae under water-deficit stress, with a focus on several key research objectives: 1) Quantify root length colonization of A. delicata in different wheat cultivars under well-watered and water-deficit conditions. 2) Assess the effects of A. delicata symbiosis on key life history traits of S. avenae , including developmental duration, longevity, fecundity, and adult weight, across water regimes and cultivars. 3) Measure aphid honeydew production and evaluate changes in aphid fresh and dry mass associated with A. delicata under drought stress. 4) Determine aphid host choice preference between drought-resistant and drought-susceptible wheat cultivars with and without A. delicata symbiosis. 5) Evaluate the impact of A. delicata on wheat root and shoot biomass under aphid infestation in both well-watered and water-deficit environments. Materials and methods 2.1 Aphids and plants The anholocyclic S. avenae clones were collected near the campus of Northwest A&F University. These clones were then reared on the Aikang-58 variety of wheat under environmental control condition (16L: 8D; 22 ± 2°C). The seeds of wheat cultivars Yunhao-618 and Xinong-1376 were used in this study. Seeds were sown in plastic cups, which contained a mixture of turfy soil, vermiculite, and perlite in a 4:3:1 volumetric ratio. This mixture offers several benefits for plant growth (Supreme 2024 ). The plastic cups were covered with transparent plastic tubes that were 7 cm in diameter and 20 cm in height. These tubes had a net on top with a 60-mesh size for ventilation purposes. This setup ensured optimal growth conditions by balancing moisture retention, drainage, and aeration, coupled with adequate light exposure and ventilation. 2.2 Management of water conditions In this study, we managed two water levels well-watered and water-deficit stress, as described previously in Dai et al. ( 2015 ). Concisely, wheat seedlings were treated with 21 ml and 7 ml of water, respectively, every 4–5 days. The seedlings were grown in 250 g (air-dry weight) of a growing medium consisting of a mixture of autoclaved quartz sand and soil inoculated with A. delicata . For the non-association control, experimental cups did not contain the AMF A. delicata inoculum with the growing media (a mixture of quartz-sand and soil). Required water conditions were maintained by measuring soil water potential and weighing the plants with the growing medium weekly by following the protocols described in Liu et al. ( 2018a ). Small stainless-steel pans were placed at the base of all cups to deliver an appropriate volume of water as needed. 2.3 Propagation of the arbuscular mycorrhizal fungus The AMF Acaulospora delicata (BGCBJ02B) was obtained from the College of Natural Resources and Environmental, Northwest A&F University, China. For propagation, local maize ( Zea mays ) was used as the host plant. The selected propagation medium consisted of 25% soil and 75% quartz sand, ensuring an optimal environment for AMF development. To ensure a contamination-free environment, the surface of 25 maize seeds was sterilized using a combination of sodium hypochlorite and ethanol. The given method for sterilizing maize seeds follows to established protocols, involving a 70% ethanol soak for 2–3 minutes followed by a 5% sodium hypochlorite soak for 5 minutes, followed by thorough rinsing with sterile water, to prevent contamination in subsequent stages of fungal inoculation and plant growth. This multi-step sterilization process helps to eliminate any potential pathogens while maintaining seed viability. Post-sterilization, the seeds were placed on a moist filter paper within a Petri dish (150 mm in diameter). The moisture levels in the Petri dish were meticulously maintained until germination occurred, ensuring that the seeds had adequate water for successful sprouting. An 8-liter plastic container (dimensions: 27×20×17 cm) was used to house the pre-germinated maize seeds. The container was filled with a growth medium composed of 8 kilograms of pasteurized loam and sand in a 1:3 volumetric ratios. Pasteurization of the medium is essential to eliminate any unwanted organisms that could interfere with AMF propagation. On top of the growth medium, 500 grams of A. delicata fungal inoculum containing spores and root fragments was added. This inoculum was thoroughly mixed with the top layer of the medium to ensure even distribution. To complete the setup, an additional 500 grams of the sterilized medium was added on top, covering the inoculum and providing a protective layer. The pre-germinated maize seeds were carefully placed on the prepared medium in the 8-liter plastic containers. The mixture of pasteurized loam and sand, combined with the A. delicata inoculum, provides a conducive environment for both the host plant and the fungus. Additionally, the layering technique ensures that the AMF inoculum is well-integrated within the medium to promote symbiosis between the maize roots and the mycorrhizal fungus (Parnell et al., 2024 ). The containers were placed in a climate chamber that provided controlled environmental conditions for four months under the following conditions: 16:8-hour light/dark cycle was employed to simulate natural light conditions, the temperature in the climate chamber was maintained at a range of 22 − 20℃, and the relative humidity was kept constant at 65% (Telli et al. 2020 ). The plants were watered with the Hoagland nutrient solution described by Hoagland and Snyder ( 1934 ), once a week. The colonization of maize roots by the arbuscular mycorrhizal fungus A. delicata was evaluated. The presence of mycorrhizal spores was confirmed by using the sieving method as described by Gerdemann and Nicholson ( 1963 ). This method involves filtering the soil sample through filter paper, followed by transferring the filtered material to a Petri dish where the number of spores is counted. This technique is essential for accurately assessing the spore presence, thereby confirming mycorrhizal colonization levels in the root samples. 2.4 Arbuscular mycorrhizal inoculation of wheat plants Drought-resistant (Yunhao-618) and drought non-resistant (Xinong-1376) cultivars of wheat were used in this experiment. The surface seed sterilization process was carried out using the method described above. After surface sterilization, both cultivars were placed on clean and moist filter paper in the petri dish (150 mm in diameter) under controlled environmental conditions of 22 ± 2℃ (Zhang et al. 2024 ). The filter paper was moistened with distilled water, and the petri plates were sealed with parafilm to maintain moisture levels. Upon successful germination, pre-germinated seeds were transferred to transparent plastic cups (7 cm in diameter). One pre-germinated seed was sown in each cup, ensuring careful handling to avoid damage to the emerging radicle (Rachel 2023 ). Each cup contained approximately 100 g mixture of autoclaved quartz sand and soil. The quartz sand was sterilized at 130℃ for 30 minutes using an autoclave and allowed to cool down for 12 hours before the experiment (Scott 2023 ). This ensures the elimination of microbial contaminants that could interfere with the experiment. Next, a layer of 25 g of A. delicata inoculum was added to the soil in each cup. A pre-germinated seed was then inserted on top of this layer of inoculum, followed by covering the seed with approximately 25 g of soil (Hibilik et al. 2019 ). This setup was designed to promote the establishment of an associative relationship between the wheat plant roots and the mycorrhizal fungus. At the end of the experiment, the roots were carefully extracted from the soil (Khoso et al. 2025b ). 2.5 Arbuscular mycorrhizal fungus root cleaning and assessment of fungal colonization The roots of A. delicata -inoculated wheat were initially removed from cups, thoroughly washed with regular tap water, and cut into segments 0.5-1 cm in length. These segments were then placed in 10 ml centrifuge tubes containing a weak formalin-acetic acid-alcohol (FAA) solution, composed of 50 ml formalin, 50 ml glacial acetic acid, and 900 ml ethyl alcohol, and stored at room temperature until further processing. After FAA treatment, the root segments underwent a soaking process in a 10% potassium hydroxide (KOH) solution (m/v) for 1 hour at 90°C to facilitate clearance and softening. Following the KOH treatment, the segments were rinsed with tap water before being subjected to an additional bleaching step. This involved immersion in a 10% alkaline solution at 20°C for 30 minutes, ensuring that any residual alkaline was thoroughly removed through multiple rinses. Subsequent to the bleaching process, the root segments were acidified by soaking them in a 1% hydrochloric acid (v/v) solution for a brief period of 3–5 minutes. This acidification prepared the roots for the staining procedure. The segments were then stained using a 0.01% fuchsine acid dye at 90°C for 10 minutes. Following the staining, the roots were decolorized using a lactophenol staining solution, allowing for satisfactory visibility of the fungal structures within the roots. After cleaning and staining, the root segments were prepared for the detection and assessment of A. delicata colonization following the methods described by Khoso et al. ( 2025b ). The processed root samples were taken from the centrifuge tubes and placed in glass petri dishes for de-staining and observation. Ten 1 cm-long root segments from both A. delicata symbiosis treatments and non-symbiosis controls were randomly selected and mounted on biological microscope slides with coverslips for examination. The prepared slides were examined under an Olympus microscope (Olympus Model: U-LH100-3 Olympus Corporation, Tokyo 163–0914, Japan) equipped with imaging software (cellSens Ver.1.7.1). The total root length colonization (TRLC) was estimated according to the methodology provided by Khoso et al. ( 2025b ). The intensity of A. delicata colonization on individual root segments was evaluated using a rating scale from 0–5 (Kormanik and McGraw 1982 ), with calculations performed using the Mycocalc software https://www2.dijon.inra.fr/mychintec/Mycocalc-prg/download.html ) according to Trouvelot et al. ( 1986 ), enabling precise quantification of colonization parameters. 2.6 Aphid performance bioassays Healthy S. avenae adults were first transferred onto new wheat seedlings. On the next day, single newly emerged neonate nymphs (< 24 h old) were transferred to wheat seedlings of each cultivar in plastic cups (7 cm in diameter, one nymph per cup) using a camel hair brush. These plastic cups were well covered with transparent plastic tubes (30 cm in height, 6 cm in diameter), which had a net on top (60 mesh) for ventilation. Stainless steel pans were placed at the base of all cups to provide a suitable amount of water when needed. The fecundity, developmental time, and adult body weight of S. avenae fed wheat were determined using the following steps. After pre-germination of the seeds, wheat seedlings were cultured in plastic cups filled with a medium consisting of a mixture of 25% soil and 75% quartz sand. This medium was sterilized at 130°C for 30 minutes and included 25 g of A. delicata inoculum (Pokorny 2024 ). Newly born nymphs of S. avenae were placed on 7-day old wheat plants (1st wheat leaf) in plastic cups. Teneral adults were weighed on a micro-balance (METTLER-TOLEDO, XS3DU, Greifensee, Switzerland) and then transferred back to the test plants. These adults were left undisturbed and monitored daily until all test aphid individuals died. Newly born aphids were removed every day to avoid overcrowding. To ensure reliability, 40 replicates were observed for each wheat cultivar grown under both water-deficit-stressed and well-watered conditions. 2.7 Honeydew excretion assay In this assay, we compared the mass of honeydew excreted by the aphids, S. avenae , when fed on drought-resistant Yunhao-618 and drought-non-resistant Xinong-1376, under well-watered and water deficit-stressed conditions. Wheat seeds were planted as above mentioned. One apterous S. avenae adult was restricted to the first leaf of each wheat plant using a self-made clip cage (Kou et al., 2022 ). The self-made clip cages were essential for confining aphids and ensuring accurate observation of honeydew production. After 24 hours, all young nymphs produced by the adult aphid were discarded to prevent overcrowding and ensure the accuracy of data. Following the initial setup period, a small piece of dried Jinjiayou aluminum foil (3 cm in diameter) Model: 613 (manufactured by Dongguan Juyou Paper & Plastic Products Co., Ltd, Guangdong China) was placed around the aphid-settled leaf inside each clip cage for an additional 24 hours. This setup ensured that honeydew excreted by the aphid would fall onto the aluminum foil for collection. The aluminum foil paper was dried to a constant weight in a drying oven at 60℃ for 30 minutes both before and after honeydew collection. This step was crucial to ensure that the weight of the honeydew could be accurately measured. The collected honeydew was then weighed using a microbalance, specifically the METTLER-TOLEDO XS3DU, which has a readability of 1 microgram. This precise measurement enabled an accurate comparison of honeydew excretion between the two wheat cultivars. A total of 40 replications were carried out for each wheat cultivar under both well-watered and water deficit-stressed conditions. Replication is a fundamental aspect of experimental design, ensuring the reliability and validity of the results. 2.8 Aphid host choice The experimental structure consisted of two 30 cm³ cages, connected by a 60 cm × 15 cm transparent plastic tunnel. Each trial involved the use of one plant cultivar at a time, with 10 A. delicate -inoculated plants placed in one cage and 10 non-symbiosis control plants placed in the other cage. The experimental design incorporated regular switching of the positions of the two cages to avoid positional effects, ensuring consistent environmental conditions for both treatment groups throughout the experiment (Guo et al. 2022 ). Approximately fifty alate aphids were released in the center of the transparent plastic tunnel (4 cm in diameter) for each trial. This strategic positioning in the middle of the tunnel allowed the aphids to distribute themselves freely between the two cages. The numbers of aphids that settled within each cage were recorded at specific intervals: 2, 4, 8, 24, 48, and 72 hours post-release. All aphid individuals on a plant within a cage were considered settlements. The experimental design focused on recording the number of aphids settled on plants with AMF symbiosis ( A. delicata –inoculated) and those without (non-symbiosis control). The consistent timing intervals allowed for detailed observation of aphid preferences over time, contributing to the understanding of how AMF symbiosis influences aphid host choice. Detailed comparative data was collected for each time point to analyze trends and patterns in aphid behavior. To ensure the reliability and robustness of the results, the experiment was conducted with a total of 20 replicates for each treatment. This high level of replication provided a robust dataset, allowing for statistical analysis to deduce significant differences and patterns in aphid settlement behavior between the two wheat cultivars. 2.9 Aphid desiccation assay Using the above-mentioned method, aphid individuals of S. avenae were raised on drought-resistant (Yunhao-618) and drought-non-resistant (Xinong-1376) wheat cultivars under well-watered and water deficit stressed levels for the bioassays of desiccation traits. A transparent desiccator glass pot, measuring 12" ×16", was used to maintain S. avenae individuals under desiccation stress. The desiccator is an airtight chamber designed to regulate humidity levels effectively, making it ideal for experiments requiring controlled moisture conditions. The pot was carefully covered with a lid to ensure an airtight seal, which is essential to prevent atmospheric moisture from entering the chamber. To maintain the desired relative humidity within the desiccator, a layer of silica gel was used. Non-toxic and suitable silica gel, no risk of chemical interaction (purchased from Shandong Rushan Taihe Co., Ltd.), to help maintain the desired relative humidity. The desiccator pot was equipped with a digital hygrothermograph recorder (Anymeter, TH20, Guangzhou, Guangdong, China), to provide real-time monitoring of temperature and humidity. For each trial, a total of ten aphid individuals were put into a 10 ml centrifuge tube. The centrifuge tube was modified to create a hole and had a net (60 mesh) to cover the hole for ventilation. This setup ensured adequate airflow and prevented the aphids from escaping. Test aphid individuals were maintained under a specific environmental condition, which in this case was 10% relative humidity (RH), for about 12 hours. During this period, the survival rates of the aphids were meticulously recorded to assess the impact of low humidity on their viability (Yang et al. 2022 ). We weighed the test S. avenae adults on a microbalance (METTLER-TOLEDO, XS3DU, Greifensee, Switzerland) to determine their fresh body mass (Mf). After they were subjected to the desiccation stress treatment for 12 h in a desiccator, test aphid individuals were reweighed to obtain the body mass after the desiccation exposure of 12 h (M 12 h). Thus, the rate of water loss per hour in this study refers to the mean water loss during the initial 12 hours of desiccation stress. After they were dead, test aphid individuals were dried at 60℃ for 60 h and then weighed again to obtain the dry mass (Md). We estimated the water loss rates using the exponential model M 12 h = Mf e- kt , and calculated the total water loss rates as kt = -ln (M 12 h/M f ) (Wharton 1985 ). Following Gibbs et al. ( 1997 ), absolute water contents were calculated as M f - M d and relative water contents as (M f- M d)/ M f ×100). 2.10 Plant developmental traits AM fungus inoculation occurred at the time of sowing seeds. First, cups of 4 cm diameter were filled with approximately 175 g of sand. Then 50 g of inoculum was mixed with the top layer of the soil, and a pre-germinated seed was added on top of the inoculum-mixed soil, which was covered on top with approximately 25 g medium (sand and soil). The control (i.e., non-symbiosis) plants were treated identically but used inoculum that had been autoclaved and allowed to dry for 24 hours before sowing. All the treated wheat plants were kept in a controlled environmental room (21 ± 2℃, relative humidity of 65 ± 5%), and the cups were placed in a randomized block design with a total of 40 replications for each treatment. Plant heights were measured, and numbers of leaves were counted weekly. At the end of the experiment, all plant samples were taken from the cups. Before drying the plant material, fresh and dry weights of whole plants were obtained by using an electronic analytical balance (Sartorius TE20101-L). The shoots, stems and leaves were oven-dried at 75℃ for 60h. Ten replicates were performed for each treatment. 3.11 Statistical analyses A randomized block (i.e., batch) design was used for the experiments. The bioassays were conducted in two batches (five replicates per treatment in each batch), with test aphid individuals randomly assigned to the treatments (i.e., symbiosis, water level and cultivar) within each batch. Ten replicates were conducted for each treatment. The data on mycorrhizal root colonization were compared with two-way ANOVA, and the effects of independent factors (i.e., cultivar and water level) were analyzed, as well as the interaction between cultivar and water level. Similarly, the total amino acid contents of wheat plants were examined with two-way ANOVA. We used three-way ANOVA to analyze the data on the developmental durations of nymphal instars, adult weight, fecundity, plant development parameters (i.e., fresh and dry weight), honeydew weight, and aphid desiccation as response variables, and the effects of independent factors, including symbiosis, water level and cultivar, were analyzed, as well as the interactions among the three factors. Mean separations were done with Tukey HSD (honestly significant differences) tests after significant ANOVA. We used the independent sample student’s t -test to identify significant differences between the association treatment and the non-association control ( P < 0.05) for settlement of S. avenae over the course of 72 hours. Data were analyzed by using Statistix 8.1, and all figures were generated by using the OriginPro 2024 software. Results 3.1 AMF A. delicata percentage root length colonization (%RLC) AMF A. delicata RLC was observed in drought resistant (Yunhan-618) and drought resistant (Xinnong-1376) wheat cultivars under different water conditions. For RLC, it was found that water level has a significant impact (Fig. 1 ; F = 334.47; df = 1.72; P = 0.001). Under water deficit-stress, high percentage of AMF A. delicata spore population was recorded in Xinong-1376 and Yunhao-618 cultivars with 67% and 62%, respectively. While control plants did not show any associations with spore production. Xinong-1376 and Yunhao-618 plants may form beneficial associations with AMF, which significantly enhances their drought tolerance. The colonization A. delicata was clearly shown in the fungus inoculated wheat plants (Fig. 2 A), as compared to control plants exhibiting zero colonization (Fig. 2 B). 3.2 Life history traits of S. avenae aphids under the influence of mycorrhizal fungus A. delicata associations 3.2.1 Developmental duration Developmental duration (DD) of S. avenae aphid, the research shows significant effects of various factors on its DD, including cultivars, water levels, and mycorrhizal fungus A. delıcata associations. The effects of association on first instar nymphs that the DD for the first instar nymphs of S. avenae prolonged with mycorrhizal non-associations on Xinong-1376 plants under water-deficit conditions. Conversely, these nymphs exhibit shorter under well-watered conditions on the Yunhao-618 cultivar. For DDs of first instar nymphs, there were no significant interactions between test factors identified. The significant effects of these conditions (Fig. 3 A; cultivars: F = 7.14; df = 1, 72; P < 0.05; associations: F = 12.21; df = 1, 72; P < 0.001; and water levels: F = 18.63; df = 1, 72; P < 0.001) for aphid development in varying moisture environments. The DD of second instar S. avenae nymphs was significantly affected by AMF A. delicata association. Significant interactions were found between mycorrhizal association and cultivars (Fig. 3 B; F = 14.58; df = 1, 72; P < 0.001); and their interactions between test factors ( F = 6.48; df = 1, 72; P = 0.05) identified. The association with AMF A. delicata significantly affected the development duration of second instar nymphs ( F = 8.82; df = 1, 72; P < 0.05). For the DD third instar nymphs of S. avenae , there were significant effects of different cultivars (Fig. 3 C; F = 9.42; df = 1, 72; P < 0.05). For DD of fourth instar nymphs, there were no significant interactions between test factors identified. The association treatments showed longer DD when associated with mycorrhizal fungi on Yunhao-618 plants under well-watered conditions. In contrast, the DD on Xinong-1376 under water deficit stressed conditions was notably shorter (Fig. 3 D). Significant interactions between cultivar and association were identified for the total developmental time of S. avenae nymphs (Fig. 4 ; F = 5.83; df = 1, 72; P < 0.05), and cultivar and water levels ( F = 4.00; df = 1, 72; P < 0.05). The total DD for S. avenae under well-watered conditions was longer (9.5 days) on mycorrhizal non-associated Xinong-1376 plants compared to a shorter DD (7 days) under water deficit-stressed conditions. 3.2.2 Aphid longevity The longevity of S. avenae on plants without A. delicata association was shorter compared to those with the association across both cultivars and water treatments. It was significant interactions between association and cultivar (Fig. 5 ; F = 8.96; df = 1, 72; P < 0.05), and water level ( F = 85.52; df = 1, 72; P < 0.001), but no significant interactions were found for this aphid. For example, under water-deficit conditions, the longevity of S. avenae on Xinong-1376 was significantly shorter for the A. delicata non-association control (mean = 13.8 d) compared to the association treatment (mean = 24.9 d). Similarly, under well-watered conditions, S. avenae on Yunhao-618 had shorter longevity in the non-association control (mean = 14.7 d) compared to the association treatment (under well-watered: mean = 21.8 d; and under water deficit-stressed: mean = 21.6 d). 3.2.3 10-d fecundity The presence of AMF A. delicata appears to beneficially affect S. avenae fecundity (Fig. 6 ; F = 10.73, df = 1, 72; P < 0.001). Water conditions also play a crucial role in determining aphid fecundity ( F = 151.37, df = 1, 72; P < 0.001). However, the association with A. delicata seems to mitigate some of these negative effects, as indicated by the higher mean fecundity under water-deficit stress in plants associated with A. delicata (mean = 22.4, SE = 1.00) compared to non-associated plants (mean = 10.2, SE = 0.85). The cultivar Yunhao-618 showed higher fecundities in association with A. delicata compared to Xinong-1376. This indicates that genetic differences between cultivars can influence the effectiveness of the A. delicata association in enhancing S. avenae fecundity (interaction between cultivar and association: F = 14.94; df = 1, 72; P < 0.001). 3.2.4 Adult weight In the context of AMF A. delicata association treatments, S. avenae exhibited higher adult weights on both Yunhao-618 and Xinong-1376 cultivars under well-watered conditions. There were significant differences due to the association (Fig. 7 ; F = 27.23; df = 1, 72; P < 0.001) and water level ( F = 85.44; df = 1, 22; P < 0.001) factors. When comparing the performance of aphids on different cultivars, Xinnong-1376 showed higher adult body weight under water stress conditions, Specifically, the average adult weight of S. avenae on Yunhao-618 under well-watered conditions was significantly higher in the association treatment (mean = 590 µg, SE = 8.59) than in the non-association control (mean = 362.6 µg, SE = 8.93). Similarly, under water-deficit stress, the teneral adult body weight on Xinong-1376 was higher in the association treatment (mean = 665.40 µg, SE = 11.88) than in the non-association control (mean = 417.00 µg, SE = 18.9). The association treatments led to enhanced adult weights under both well-watered and water deficit-stressed conditions, which highlights the beneficial role of symbiotic relationships in mitigating the adverse effects of water stress on aphid performance. 3.3 Aphid honeydew production For honeydew production of this aphid, cultivars (Fig. 8 ; F = 14.89; df = 1,72; P < 0.001), water level ( F = 46.11; df = 1,72; P < 0.001), and for association ( F = 22.05; df = 1,72; P 0.05), or for cultivars and association ( F = 0.05; df = 1,72; P > 0.05). The amount of honeydew production by S. avenae under well-watered conditions was significantly higher than under water deficit-stressed conditions on all test wheat cultivars. With A. delicate association under both water levels, this aphid produced higher amounts of honeydew (mean = 4.27 µg, SE = 0.11) on drought non-resistant Xinong-1376 than on drought-resistant cultivars. There was lowest amounts of honeydew (mean = 0.97 µg, SE = 0.14) on drought resistant Yunhao-618 under water deficit-stressed conditions. 3.4 Aphid desiccation assays 3.4.1 Fresh mass In terms of fresh body mass, we found significant effects for cultivars (Fig. 9 A; F = 25.49; df = 1, 72; P < 0.001), water level ( F = 20.93; df = 1, 72; P < 0.001), and interactions between cultivars and association ( F = 6.78; df = 1, 72; P < 0.05). The fresh body masses of S. avneae on the drought resistant Yunhao-618 tended to be higher with A. delicate association than those on Xinong-1376 test drought non-resistant cultivars. Under water-deficit stress, high fresh body masses (mean = 375.77 µg, SE = 21.65) on Yunhao-618 and without A. delicate association lower (mean = 197.80 µg, SE = 5.15) on Xinong-1376, but in terms of A. delicate association alone, this pattern was found opposite patterns on both cultivars. 3.4.2 Dry mass For the dry mass of this aphid, we identified significant effects for cultivars (Fig. 9 B; F = 44.01; df = 1, 72; P < 0.001), and water treatments ( F = 12.70; df = 1, 72; P < 0.001), or for association ( F = 6.06; df = 1, 72; P < 0.05). On Yunhao-618, the dry body masses of this aphid tended to increase with increasing water-deficit stress, but this pattern was not found on other test cultivar (Xinong-1376). Under well-watered conditions, this aphid without A. delicate association had higher dry body masses on Yunhao-618 than on Xinong-1376 cultivars. 3.4.3 Water balance traits We found effects on absolute water contents (AWC) of S. avenae on cultivars (Fig. 10 A; F = 14.64; df = 1, 72; P < 0.001), water level ( F = 19.28; df = 1, 72; P < 0.001), and the interaction between cultivars and water level ( F = 6.60; df = 1, 72; P < 0.05), or interaction between cultivars and association ( F = 6.60; df = 1, 72; P < 0.05). The highest absolute water contents of this aphid were found with A. delicate association on drought-resistant Yunhao-618 under water-deficit stressed conditions, but this aphid under water-deficit stressed had the lowest absolute water contents on Xinong-1376. Relative water contents (RWC) of S. avenae also showed significant variability under different conditions. There were significant effects identified for cultivars (Fig. 10 B; F = 24.04; df = 1, 72; P < 0.001), association ( F = 11.23; df = 1, 72; P < 0.001), and the interaction between cultivars and water level ( F = 26.54; df = 1, 72; P < 0.001). Additionally, significant interactions across all factors ( F = 4.52; df = 1, 72; P < 0.05) were observed. RWC of S. avenae without A. delicate association on drought resistant Yunhao-618 showed a decreasing trend with increasing well-water. Oppositely, this pattern was found on decreasing trend with increasing water deficit-stress on Xinong-1376 cultivar. This aphid had equal relative water contents on both cultivars with A. delicate association under water deficit-stressed and well-watered conditions. Water loss rates (WLR) in S. avenae associated with A. delicate showed significant interactions between cultivars and water level (Fig. 10 C; F = 11.64; df = 1, 72; P < 0.001), as well as between cultivars and association ( F = 4.16; df = 1, 72; P < 0.05). Significant interactions between all factors were also noted ( F = 20.90; df = 1, 72; P < 0.001). Water loss rates of S. avenae with A. delicate association showed a decreasing trend with increasing water-deficit stress on Yunhao-618, and without association on Xinong-1376. Under well-watered, this aphid had the lowest water loss rates with A. delicate association on Yunhao-618, and Xinong-1376 under water-deficit stressed. 3.5 Aphid host choice The host choice performance of S. avenae on different wheat cultivars under varying water conditions reveals complex dynamics influenced by the plant's drought resistance and association with AMF ( A. delicate ). Under water-deficit stress, S. avenae showed a significant preference for plants associated with A. delicate over the non-association control on Yunhao-618 at 12 hours post-release (hpr) (Fig. 11 A; t = 7.40; df = 18; P < 0.001) which was also observed under well-watered conditions (Fig. 11 B; t = 2.86; df = 18; P < 0.05). A notable preference shift was seen in the Xinong-1376 cultivar under severe-water deficit at just 2 hpr (Fig. 11 C; t = 4.96; df = 18; P < 0.001) and at 12 hpr under well-watered conditions (Fig. 11 D; t = 3.89, df = 18; P < 0.001). At 72 hpr, the aphid population with A. delicate on Xinnong-1376 (under water-deficit stress mean: 19.20, SE = 0.39; and under well-watered mean: 23.50, SE = 0.23) was significantly higher than that of Yunhao-618 (under water-deficit stress mean: 5.50, SE = 0.11; and under well-watered mean: 9.30, SE = 0.26). At 72 hpr, S. avenae showed with A. delicate a strong preference for the drought non-resistant Xinong-1376 over the drought-resistant Yunhao-618 cultivar. It can be partly explained by the sustainability and nutrition provided by the respective plants under both water-deficit and well-watered conditions with A. delicate . The mean aphid population was significantly higher on Xinong-1376. 3.6 Plant developmental traits The study found that wheat plants with AMF ( A. delicate ) association treatments had significantly larger fresh weights compared to those without the associations (Fig. 12 A; association: F = 40.05; df = 1, 72; P < 0.001). Especially, under conditions of well-watered condition, the fresh plant weight of AMF combination is higher compared to water-deficit stressed conditions (water level: F = 29.76; df = 1,72; P < 0.001). The effect of plant dry weight on cultivars (Fig. 12 B; F = 5.56༛ df = 1,72༛ P < 0.05), water level ( F = 11.80; df = 1,72; P < 0.001) and association ( F = 5.20; df = 1.72; P < 0.05) had a significant influence. In addition, the interaction between associations and cultivar ( F = 5.56; df = 1, 72; P < 0.05) was significant. Discussion The high root length colonization (%RLC) by AMF A. delicata observed in drought-resistant wheat cultivars Xinong-1376 (67%) and Yunhao-618 (62%) under water deficit stress underscores the important role of AMF in enhancing drought tolerance. This aligns with recent studies demonstrating that effective AMF colonization improves water and nutrient uptake, crucial for maintaining plant physiological functions during drought (Nopphakat et al. 2022; Akensous et al. 2024 ). Similar findings show that AMF symbiosis increases root surface area and spore production, facilitating access to deeper soil moisture (Kang et al. 2022 ; Li et al. 2025 ). Furthermore, research highlights that AMF-mediated enhancements in antioxidant enzyme activity reduce oxidative stress under drought, contributing to improved plant resilience (Li et al. 2025 ). Importantly, AMF colonization is cultivar-specific, influenced by root architecture and genetic compatibility, as observed in Xinong-1376 and Yunhao-618, suggesting targeted AMF inoculation could optimize drought adaptation (Duan et al. 2024 ). These synergistic insights affirm AMF A. delicata as a promising bio-fertilizer to bolster wheat productivity in water-limited environments. The DD of S. avenae nymphs was notably influenced by the combination of wheat cultivars, water availability, and association with AMF, particularly A. delicata . Under water-deficit conditions, first instar nymphs showed prolonged DD on Xinong-1376 plants without AMF, indicating that drought stress and the absence of beneficial fungal symbiosis negatively impact aphid development (Babikova et al. 2014 ; Liu et al. 2018b ; Stallmann and Schweiger 2021). In contrast, under well-watered conditions, Yunhao-618 plants with AMF supported faster development, suggesting that AMF may mitigate stress under optimal water availability (Thompson, 2022 ). The second instar nymphs exhibited longer DD on Xinong-1376 without AMF under well-watered conditions, while under drought, DD shortened—suggesting stress may accelerate development in the absence of AMF. Similar findings by Kula et al. ( 2005 ) and Cascone et al. ( 2024 ) show that AMF often reduce insect development time under optimal conditions by improving plant nutrition, but under stress, absence of AMF can lead to rapid aphid development. Cultivar selection also influenced aphid DD, though no significant interaction between cultivar and other factors was observed (Platkova et al. 2020 ). For third instar nymphs, A. delicata shortened DD on Xinong-1376 under both water regimes, while under drought stress, Yunhao-618 without AMF caused significant DD prolongation. This suggests AMF can alleviate drought effects and enhance aphid development (Yang et al. 2014 ; Liu et al. 2018a ). For fourth instars, longer DD was observed on Yunhao-618 with AMF under well-watered conditions, possibly due to AMF-induced plant defenses (Babikova et al. 2013 ). Conversely, shorter DD under drought on Xinong-1376 supports the idea that aphids may accelerate development to compensate for reduced plant quality (Hodge and Storer 2015 ; Gange et al. 2019 ). Overall, aphid development is context-dependent, shaped by the complex interactions among plant genotype, AMF association, and water availability, consistent with findings that water stress and AMF play key roles in modulating aphid fitness and host plant quality (Beetge and Krüger 2019 ). Significant interactions between wheat cultivar and mycorrhizal association, as well as between cultivar and water levels, strongly influence the total developmental duration (DD) of S. avenae nymphs. The longer DD (9.5 days) observed on mycorrhizal non-associated Xinong-1376 plants under well-watered conditions compared to the shorter DD (7 days) under water deficit stress aligns with recent studies showing that optimal water availability promotes aphid development by enhancing plant vigor and nutrient quality. Conversely, drought stress accelerates aphid development, possibly as a compensatory adaptation to reduced host quality. Similar findings by Liu et al. (2018) indicated that drought stress shortens aphid developmental periods, especially in the absence of beneficial fungal symbiosis. Moreover, AMF associations can moderate these effects by improving plant nutrition and stress tolerance, sometimes prolonging aphid development under favorable conditions (Babikova et al. 2014 ). The observed increase in S. avenae fecundity on wheat plants colonized by A. delicata under water-deficit stress aligns with emerging research demonstrating that AMF can enhance host plant nutritional and physiological status, thereby benefiting phloem-feeding herbivores. Similar studies highlight AMF’s role in improving plant water and nutrient uptake under drought, which maintains or elevates phloem sap quality crucial for aphid reproduction (Gange 2007 ; Zeng 2024 ). The cultivar-specific response, where Yunhao-618 shows heightened aphid fecundity under AMF symbiosis, underscores genetic variation in plant-mycorrhizal compatibility affecting herbivore performance (Khoso et al. 2025b ). Contrastingly, research involving fungal endosymbionts in other aphid species found reduced fecundity and lifespan, suggesting that microbial associations can have positive or negative effects on herbivores depending on symbiont type and plant context (Meister et al. 2006 ; Johnson et al. 2011 ). Additionally, drought typically reduces aphid fecundity and population growth (Kansman et al. 2022 ), but the buffering effect of A. delicata suggests potential for leveraging AMF in integrated pest and drought management strategies to sustain crop resilience and manage aphid outbreaks effectively. Our findings align with recent research highlighting the role of both wheat cultivar traits and AMF in shaping aphid performance under varying water conditions. Consistent with Liu et al. ( 2018a ), differences in adult S. avenae weight across water treatments reflect the aphid’s adaptation potential, influenced by plant nutritional quality and stress resilience. AMF-driven enhancements in nutrient and water uptake improve host plant vigor, creating a more favorable environment for aphid growth, as seen in the higher adult weights on Xinong-1376 compared to Yunhao-618 under drought stress. This supports previous work showing drought-resilient cultivars maintain better aphid nutritional support (Pineda et al. 2017 ). However, severe water deficit generally reduces aphid fitness due to diminished plant resources, consistent with Stiling et al. ( 2009 ). The positive influence of A. delicata on aphid weight under both water regimes indicates AMF can buffer drought effects while simultaneously promoting aphid development, a dynamic critical for predicting pest outbreaks under climate change (Rizzo 2017 ). This interplay poses challenges for crop protection as increased aphid vigor may lead to more severe yield losses (Lenoir et al. 2016 ; CABI 2021 ). Aphid honeydew production varies significantly with host plant cultivar, water availability, and AMF associations, consistent with findings from previous research. Well-watered plants generally support higher honeydew output due to increased nutrient availability and more efficient aphid feeding, aligning with Huberty and Denno ( 2004 ) and van Rooijen ( 2023 ). Plant stress, such as drought, reduces phloem pressure and increases sap viscosity, hindering aphid feeding and lowering honeydew production, as observed by Kansman et al. ( 2020 ). Cultivar differences further influence these dynamics; drought-susceptible cultivars like Xinong-1376 provide more favorable feeding conditions for aphids regardless of water regime, leading to higher honeydew outputs. In contrast, resistant cultivars such as Yunhao-618 exhibit physical and chemical defenses that reduce aphid feeding efficiency and honeydew production under drought stress (Stallmann et al. 2022; Tous-Fandos et al. 2023 ). AMF association with A. delicata enhances honeydew production, likely by improving phloem sap access or modulating plant defenses, supporting concepts from Styrsky and Eubanks ( 2007 ) and Milcu et al. ( 2015 ). These insights emphasize the complex, genotype-environment-microbe interplay regulating aphid-plant interactions and pest dynamics under stress. The observed influence of A. delicata symbiosis on aphid fresh and dry body mass under drought conditions echoes recent findings on the integral role of microbial associations in aphid physiology and adaptation. Similar to Csorba et al. ( 2024 ), symbiotic relationships with microbes such as Buchnera aphidicola enhance aphid nutrient acquisition, critical under host plant stress. The cultivar-specific benefits, with Yunhao-618 facilitating higher aphid mass under drought, reflect differential plant physiological responses consistent with Pozo and Azcón-Aguilar ( 2007 ), who emphasize that drought-resistant cultivars better exploit AM fungi symbiosis to maintain plant health. Recent research also links increased aphid performance on drought-stressed resistant cultivars to altered phloem chemistry, such as elevated soluble sugars and nitrogen availability, supporting Huberty and Denno’s ( 2004 ) and Kansman et al.’s ( 2022 ) conclusions. Conversely, susceptible cultivars like Xinong-1376 experience compromised physiology under drought, reducing aphid support. The findings on the absolute water content (AWC), relative water content (RWC), and water loss rate (WLR) of S. avenae reveal intricate interactions between aphids, host plant cultivars, water availability, and symbiotic association with A. delicata . Echoing recent research on other herbivores such as Aphis nerii and leaf-cutting ants (Carvajal Acosta et al. 2023 ; Gely et al. 2020 ), plant genotype and environmental stressors play pivotal roles in determining insect water status and physiological performance. The higher AWC and maintained RWC of S. avenae on the drought-resistant cultivar Yunhao-618 under water deficit, especially when associated with AMF, align with studies demonstrating that mycorrhizal fungi enhance host plant water retention, nutrient uptake, and overall physiological resilience under drought (Díaz-Hernández et al. 2024 ; Yan et al. 2022 ). These improvements indirectly benefit associated herbivores by stabilizing phloem sap quality and water availability. In contrast, drought-susceptible cultivars like Xinong-1376 show reduced aphid water content and higher water loss rates under stress, mirroring findings in other insect-plant systems where plant stress diminishes herbivore hydration and fitness (Benoit et al. 2023 ). The mitigation of water loss in S. avenae by AMF symbiosis suggests symbiotic fungi may contribute to improved insect desiccation tolerance, a trait critical under climate change-induced drought scenarios (Addo-Bediako et al. 2001 ). Moreover, the persistent aphid preference for AMF-associated plants across water regimes supports the hypothesis that mycorrhizal fungi improve plant nutritional quality and attractiveness independent of water status, corroborated by findings in other plant-insect studies (Charters et al. 2022 ; Bell et al. 2024 ). Overall, this body of research highlights a complex, multi-layered network where plant genotype, microbial symbiosis, and environmental stress interact to shape herbivore physiology, behavior, and survival. Understanding these dynamics is essential for developing sustainable pest and water management strategies in agroecosystems facing increasing water scarcity. The preference of S. avenae for drought-sensitive wheat cultivars like Xinong-1376 over drought-resistant ones such as Yunhao-618, especially when associated with A. delicata , aligns with recent findings on plant–aphid–AMF interactions under water stress. Similar research highlights that drought-stressed, less resistant cultivars often accumulate higher concentrations of soluble nitrogen compounds and amino acids, improving phloem sap quality and thus aphid nutrition and reproduction (Weldegergis et al. 2015 ; Kansman et al. 2020 ). The temporal dynamics in aphid colonization, with fluctuations corresponding to plant defense activation and subsequent weakening, reflect complex physiological responses modulated by AMF symbiosis, mirroring observations in other studies where initial plant stress induces defensive compounds that later decline, increasing herbivore susceptibility (Kansman et al. 2022 ). Furthermore, research on AMF-mediated drought tolerance shows that while mycorrhizal association can enhance plant resistance, cultivar-specific variability often determines the degree to which aphid colonization is suppressed or facilitated (Volpe et al. 2018 ; Sonbol et al. 2025 ). The observed increase in dry and fresh weights of wheat cultivars Xinong-1376 and Yunhao-618 associated with A. delicata under both well-watered and drought conditions aligns well with recent findings on AMF-mediated plant growth promotion. Similar studies across diverse crops have consistently demonstrated AMF’s role in enhancing biomass by improving nutrient and water uptake through an extensive hyphal network, especially under abiotic stress such as drought (Gao et al. 2023 ). For instance, AMF inoculation in tomatoes and cucumbers significantly increased fresh and dry weights, attributed to enhanced photosynthetic efficiency and antioxidant defense that mitigate oxidative stress during water deficit (Alam et al. 2023 ; Tang et al. 2022 ). These mechanisms contribute to stabilization of water potential and accumulation of protective metabolites like flavonoids and glycosides (Stallmann et al. 2020). Moreover, cultivar-specific responses to AMF are increasingly recognized, with drought-resistant varieties like Yunhao-618 leveraging AMF symbiosis to sustain growth under stress, consistent with findings in barley and maize (Begum et al. 2019 ). Additionally, enhanced root biomass observed with AMF inoculation supports stronger water and nutrient acquisition capacity even in varied drought intensities (Chandrasekaran 2022 ). This reinforces the potential of integrating AMF inoculation into crop management strategies to improve yield stability under fluctuating water availability. Collectively, these studies underscore AMF’s broad applicability and cultivar-specific benefits for sustainable agriculture under climate variability. Conclusion This study revealed that A. delicata colonization (62–67% RLC) significantly enhanced Sitobion avenae performance on both drought-resistant and drought-susceptible wheat cultivars under contrasting water regimes. AMF symbiosis prolonged aphid longevity (13.8 to 24.9 days), increased fecundity (10.2 to 22.4 offspring), and elevated adult body weight (362.6 to 665.40 µg) under water-deficit stress. Enhanced aphid desiccation tolerance was evident through increased fresh body mass, absolute water contents, and reduced water loss rates on AMF-colonized plants. Host preference experiments demonstrated strong aphid attraction to mycorrhizal-associated plants, with population densities reaching 19.20–23.50 individuals on Xinong-1376 compared to 5.50–9.30 on Yunhao-618 at 72 hours post-release. AMF colonization significantly improved plant biomass production under both water treatments, confirming beneficial effects on host plant fitness. These results indicate that mycorrhizal-mediated alterations in plant nutritional quality create more suitable conditions for aphid herbivory, potentially offsetting cultivar resistance mechanisms. The tri-trophic interactions observed highlight complex ecological trade-offs where plant-beneficial symbionts may inadvertently enhance pest performance through bottom-up nutritional effects. Cultivar-specific responses suggest genetic variation influences the magnitude of AMF-aphid interactions under environmental stress. These findings emphasize the need to consider belowground symbiotic associations when developing drought-tolerant crop varieties for sustainable pest management. Future research should elucidate the biochemical mechanisms underlying these context-dependent plant-microbe-insect interactions in agricultural ecosystems. Declarations Competing interests: The author declares that there are no competing interests. Funding: The authors did not receive support from any organization for the submitted work. Author Contributions: AGK : Conceptualization, Data curation, Formal analysis, Methodology, Writing – original draft. MA : Writing – review & editing. Acknowledgments: We would like to thank Prof. Yajun Xi from College of Agronomy of Northwest A&F University for providing seeds of different wheat cultivars used in this study. 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Pergamon, Oxford, UK, pp 565–603 Yahya M, Saeed NA, Nadeem S, Hamed M, Shokat S (2017) Role of wheat varieties and insecticide applications against aphids for better wheat crop harvest. Pak J Zool 49(6):2217–2225. https://doi.org/10.17582/J.pjz/2017.49.6.2217.2225 Yan Q, Li X, Xiao X, Chen J, Liu J, Lin C, Guan R, Wang D (2022) Arbuscular mycorrhizal fungi improve the growth and drought tolerance of Cinnamomum migao by enhancing physio-biochemical responses. Ecol Evol 12(7):e9091. https://doi.org/10.1002/ece3.9091 Yang B, Wen H, Wang S, Zhang J, Wang Y, Zhang T, Yuan K, Lu L, Liu Y, Xue Q, Shan H (2024) Enhancing drought resistance and yield of wheat through inoculation with Streptomyces pactum act 12 in drought field environments. Agro 14:692. https://doi.org/10.3390/agronomy14040692 Yang H, Dai Y, Wang X, Zhang Q, Zhu L, Bian X (2014) Meta-analysis of interactions between arbuscular mycorrhizal fungi and biotic stressors of plants. Sci World J 746506. https://doi.org/10.1155/2014/746506 Yang Y, Li X, Liu D, Pei X, Khoso AG (2022) Rapid changes in composition and contents of cuticular hydrocarbons in Sitobion avenae (Hemiptera: Aphididae) clones adapting to desiccation stress. J Econ Entomol 115(2):508–518. https://doi.org/10.1093/jee/toab240 Ye Q, Wang H, Li H (2023) Arbuscular mycorrhizal fungi enhance drought stress tolerance by regulating osmotic balance, the antioxidant system, and the expression of drought-responsive genes in Vitis vinifera L. Aust J Grape Wine Res 2023(1):7208341. https://doi.org/10.1155/2023/7208341 Yooyongwech S, Phaukinsang N, Cha-um S, Supaibulwatana K (2013) Arbuscular mycorrhiza improved growth performance in Macadamia tetraphylla L. grown under water deficit stress involves soluble sugar and proline accumulation. Plant Growth Regul 69:285–293. https://doi.org/10.1007/s10725-012-9771-6 Zeng M (2024) The mutual effect of nutrients on plant-herbivore interactions. Plant Ecol 225:1035–1045. https://doi.org/10.1007/s11258-024-01452-3 Zhang R, Zhang H, Wang L, Zeng Y (2024) Effect of salt-alkali stress on seed germination of the halophyte Halostachys caspica. Sci Rep 14:13199. https://doi.org/10.1038/s41598-024-61737-5 Zhang W, Yu L, Han B, Liu K, Shao X (2022) Mycorrhizal inoculation enhances nutrient absorption and induces insect-resistant defense of Elymus nutans. Front Plant Sci 13:898969. https://doi.org/10.3389/fpls.2022.898969 Zou YN, Zhang F, Srivastava AK, Wu QS, Kuca K (2021) Arbuscular mycorrhizal fungi regulate polyamine homeostasis in roots of trifoliate orange for improved adaptation to soil moisture deficit stress. Front Plant Sci 11:600792. https://doi.org/10.3389/fpls.2020.600792 Zust T, Agrawal AA (2017) Trade-offs between plant growth and defense against insect herbivory: An emerging mechanistic synthesis. Annu Rev Plant Boil 68:513–534. https://doi.org/10.1146/annurev-arplant-042916-040856 Additional Declarations The authors declare no competing interests. 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. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. 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-8715853","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":581504708,"identity":"ad86342f-6f6c-46ea-99f1-5ab152ac2f2b","order_by":0,"name":"Abdul Ghaffar Khoso","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAAy0lEQVRIiWNgGAWjYBACgxtg8p8cAwMPkVoMZ4DIigPGxGsxlgCRZw4kNhCtxUy6+fCHj2130jccP3vwwQcGOzndBgJabGSOpUnObHuWu+FMXjLQlcnGZgcIaZHIMWPmbWPO3XAgx0yah+FA4jZCWswkcow/A7WkG5x/Q6QWY4kcA2meM4cTDG4Qa4vhjLQ0yRkVaYYzb7wxNpxhQIRfDG4kH/7wwcBGnu98juGDDxV2cgS1wIECWKUBscpBQL6BFNWjYBSMglEwogAASNJE6MgAn+MAAAAASUVORK5CYII=","orcid":"https://orcid.org/0000-0002-5224-8740","institution":"College of Plant Protection, Northwest A\u0026F University, 712100 Yangling, Shaanxi, China","correspondingAuthor":true,"prefix":"","firstName":"Abdul","middleName":"Ghaffar","lastName":"Khoso","suffix":""},{"id":581504709,"identity":"f7ad6521-8c98-4851-b646-e65fd385b42c","order_by":1,"name":"Muhammad Awais","email":"","orcid":"https://orcid.org/0000-0002-6211-405X","institution":"College of Plant Protection, Northwest A\u0026F University, 712100 Yangling, Shaanxi, China","correspondingAuthor":false,"prefix":"","firstName":"Muhammad","middleName":"","lastName":"Awais","suffix":""}],"badges":[],"createdAt":"2026-01-28 03:38:28","currentVersionCode":1,"declarations":{"humanSubjects":false,"vertebrateSubjects":true,"conflictsOfInterestStatement":false,"humanSubjectEthicalGuidelines":false,"humanSubjectConsent":false,"humanSubjectClinicalTrial":false,"humanSubjectCaseReport":false,"vertebrateSubjectEthicalGuidelines":true},"doi":"10.21203/rs.3.rs-8715853/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-8715853/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":101398547,"identity":"53c268c8-8cce-42ea-8b64-c3d47e63d1d0","added_by":"auto","created_at":"2026-01-29 09:42:13","extension":"jpeg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":74486,"visible":true,"origin":"","legend":"\u003cp\u003eMean percentage root length colonization (% RLC) of arbuscular mycorrhizal fungus \u003cem\u003eAcaulospora delicata\u003c/em\u003edrought-resistant (Yunzao-618) and drought non-resistant (Xinong-1376) seedlings under water deficit. Different letters on the bars indicate significant differences among treatments at the \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05 level, as determined by ANOVA followed by Tukey test. WDS, water-deficit stress; WW, well-watered conditions. Error bars show mean ± SE. Quantification was carried out after other experiments were completed\u003c/p\u003e","description":"","filename":"image1.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8715853/v1/b52a8f66e1080257a61ca993.jpeg"},{"id":101381453,"identity":"ffe7c278-5e08-4ea5-b0be-eaeb9a3a80d6","added_by":"auto","created_at":"2026-01-29 06:19:20","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":109755,"visible":true,"origin":"","legend":"\u003cp\u003eStructure of the arbuscular mycorrhizal (AM) fungus \u003cem\u003eAcaulospora delicata\u003c/em\u003e on colonized and non-colonized wheat seedling roots. Images were obtained under a microscope at 10x magnification after acidic fuchsin staining: (A) fungus-colonized roots; (B) non-colonized roots\u003c/p\u003e","description":"","filename":"image2.jpg","url":"https://assets-eu.researchsquare.com/files/rs-8715853/v1/40525a3585765ee8c0ce07bc.jpg"},{"id":101381455,"identity":"4f424713-ac3c-440c-96fe-b12a7b4da1c6","added_by":"auto","created_at":"2026-01-29 06:19:20","extension":"jpeg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":403635,"visible":true,"origin":"","legend":"\u003cp\u003eComparisons of the developmental times of nymphal instars (A, 1st; B, 2nd; C, 3rd; D, 4th) of \u003cem\u003eSitobion avenae\u003c/em\u003e fed on \u003cem\u003eAcaulospora delicata\u003c/em\u003e-association and non-association wheat seedlings under different water levels. Yunhao-618, drought-resistant cultivar; Xinong-1376, drought non-resistant cultivar. Different letters on the bars indicate significant differences among treatments at the \u003cem\u003eP\u003c/em\u003e\u0026lt; 0.05 level, as determined by ANOVA followed by Tukey test. WDS, water-deficit stress; WW, well-watered conditions. Error bars show mean ± SE\u003c/p\u003e","description":"","filename":"image3.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8715853/v1/77d9cc4862b5c6d713ef8610.jpeg"},{"id":101381450,"identity":"8092da98-9857-488f-987b-bd5e6e2dd92f","added_by":"auto","created_at":"2026-01-29 06:19:20","extension":"jpeg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":84604,"visible":true,"origin":"","legend":"\u003cp\u003eComparisons of the total developmental times of nymphal instars (1st-4th) for \u003cem\u003eSitobion avenae\u003c/em\u003e fed on \u003cem\u003eAcaulospora delicata\u003c/em\u003e-association and non-association wheat seedlings under different water levels. Yunhao-618, drought-resistant cultivar; Xinong-1376, drought-non-resistant cultivar. Different letters on the bars indicate significant differences among treatments at the \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05 level (ANOVA followed by Tukey test). WDS, water-deficit stress; WW, well-watered conditions. Error bars represent the mean ± SE.\u003c/p\u003e","description":"","filename":"image4.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8715853/v1/2695e5bdfe69d8541ac46a8b.jpeg"},{"id":101381459,"identity":"26427948-c160-4998-834f-b1ccb3951b2a","added_by":"auto","created_at":"2026-01-29 06:19:20","extension":"jpeg","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":74566,"visible":true,"origin":"","legend":"\u003cp\u003eComparisons of longevity for \u003cem\u003eSitobion avenae\u003c/em\u003e fed on \u003cem\u003eAcaulospora delicata\u003c/em\u003e-association and non-association wheat seedlings under different water levels. Yunhao-618, drought-resistant cultivar; Xinong-1376, drought-non-resistant cultivar. Different letters on the bars indicate significant differences among treatments at the \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05 level (ANOVA followed by Tukey test). WDS, water-deficit stress; WW, well-watered conditions. Error bars represent the mean ± SE\u003c/p\u003e","description":"","filename":"image5.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8715853/v1/6142909e738a821f5397f292.jpeg"},{"id":101751196,"identity":"a762d389-e65d-4b4a-b004-48553a6a128c","added_by":"auto","created_at":"2026-02-03 10:18:06","extension":"jpeg","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":72089,"visible":true,"origin":"","legend":"\u003cp\u003eComparisons of 10-d fecundity for \u003cem\u003eSitobion avenae\u003c/em\u003e fed on \u003cem\u003eAcaulospora delicata\u003c/em\u003e-association and non-association wheat seedlings under different water levels. Yunhao-618, drought-resistant cultivar; Xinong-1376, drought-non-resistant cultivar. Different letters on the bars indicate significant differences among treatments at the \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05 level (ANOVA followed by Tukey test). WDS, water-deficit stress; WW, well-watered conditions. Error bars represent the mean ± SE\u003c/p\u003e","description":"","filename":"image6.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8715853/v1/a2b8682d21f1eb3c7628f286.jpeg"},{"id":101381452,"identity":"11e3c387-4ebe-4313-82d0-eb4d5ac8ff1e","added_by":"auto","created_at":"2026-01-29 06:19:20","extension":"jpeg","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":80879,"visible":true,"origin":"","legend":"\u003cp\u003eComparisons of adult weight for \u003cem\u003eSitobion avenae\u003c/em\u003e fed on \u003cem\u003eAcaulospora delicata\u003c/em\u003e-association and non-association wheat seedlings under different water levels. Yunhao-618, drought-resistant cultivar; Xinong-1376 is a drought-non-resistant cultivar. Different letters on the bars indicate significant differences among treatments at the \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05 level (ANOVA followed by Tukey test). WDS, water-deficit stress; WW, well-watered conditions. Error bars represent the mean ± SE\u003c/p\u003e","description":"","filename":"image7.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8715853/v1/64903e147db4127f70a89cd0.jpeg"},{"id":101381461,"identity":"64c48146-79ea-4bea-a92d-3ec5f43ba00c","added_by":"auto","created_at":"2026-01-29 06:19:21","extension":"jpeg","order_by":8,"title":"Figure 8","display":"","copyAsset":false,"role":"figure","size":70621,"visible":true,"origin":"","legend":"\u003cp\u003eComparisons of honeydew production for \u003cem\u003eSitobion avenae\u003c/em\u003e fed on \u003cem\u003eAcaulospora delicata\u003c/em\u003e-association and non-association on drought-resistant (Yunhao-618) and drought-non-resistant (Xinong-1376) wheat cultivars under different water levels. WDS, water-deficit stress; WW, well-watered conditions. Different letters on the bars indicate significant differences among treatments at the \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05 level (ANOVA followed by Tukey test). Error bars represent the mean ± SE\u003c/p\u003e","description":"","filename":"image8.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8715853/v1/365c7305513829795a0d3363.jpeg"},{"id":101381456,"identity":"4e0e4ea8-bac5-4fca-b29d-1880f4a09e64","added_by":"auto","created_at":"2026-01-29 06:19:20","extension":"jpeg","order_by":9,"title":"Figure 9","display":"","copyAsset":false,"role":"figure","size":211019,"visible":true,"origin":"","legend":"\u003cp\u003eComparison of the body mass of \u003cem\u003eSitobion avenae\u003c/em\u003e fed on \u003cem\u003eAcaulospora delicata\u003c/em\u003e-association and non-association on drought-resistant (Yunhao-618) and drought-non-resistant (Xinong-1376) wheat cultivars under different water levels. WDS, water-deficit stress; WW, well-watered conditions. (A) Fresh body mass; (B) Dry body mass. Different letters indicate significant differences at the \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05 level, based on ANOVA followed by Tukey test. Error bars represent the mean ± SE\u003c/p\u003e","description":"","filename":"image9.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8715853/v1/5ff7e8cae48dc2b6298eadf4.jpeg"},{"id":101381462,"identity":"670934b0-2432-41c9-8045-a790ec912723","added_by":"auto","created_at":"2026-01-29 06:19:21","extension":"jpeg","order_by":10,"title":"Figure 10","display":"","copyAsset":false,"role":"figure","size":351794,"visible":true,"origin":"","legend":"\u003cp\u003eComparisons of water balance traits in \u003cem\u003eSitobion avenae\u003c/em\u003e fed on \u003cem\u003eAcaulospora delicata\u003c/em\u003e-assosiation and non-assosiation on drought-resistant (Yunhao-618) and drought-non-resistant (Xinong-1376) wheat cultivars under different water levels. WDS, water-deficit stress; WW, well-watered conditions. (A) Absolute water content; (B) Relative water content; (C) Water loss rate per hour. Different letters indicate significant differences at the \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05 level, based on ANOVA followed by Tukey test. Error bars represent the mean ± SE\u003c/p\u003e","description":"","filename":"image10.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8715853/v1/6e23afb69388e02262aa6f53.jpeg"},{"id":101398544,"identity":"8668ba13-2b04-4833-bb43-cecea2171fb2","added_by":"auto","created_at":"2026-01-29 09:42:10","extension":"jpeg","order_by":11,"title":"Figure 11","display":"","copyAsset":false,"role":"figure","size":350379,"visible":true,"origin":"","legend":"\u003cp\u003eSettlement of alate \u003cem\u003eSitobion avenae\u003c/em\u003e on \u003cem\u003eAcaulospora delicata\u003c/em\u003e-assosiation and control (non-assosiation) wheat seedlings under different water levels: (A) water-deficit stressed Yunhao-618; (B) well-watered Yunhao-618; (C) water-deficit stressed Xinong-1376; and (D) well-watered Xinong-1376. Stars above the bars indicate significant differences between treatments with and without symbiosis, identified by Student’s t-test (“*”, \u003cem\u003eP\u003c/em\u003e\u0026lt; 0.05; “\u003cstrong\u003e**\u003c/strong\u003e”,\u003cstrong\u003e \u003c/strong\u003e\u003cem\u003eP\u003c/em\u003e \u0026lt; 0.01). Error bars represent ± SE\u003c/p\u003e","description":"","filename":"image11.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8715853/v1/31502ff1e4da146555efbb1e.jpeg"},{"id":101398671,"identity":"8e7c4e04-fad3-466b-b86f-70fcb2f7062a","added_by":"auto","created_at":"2026-01-29 09:43:53","extension":"jpeg","order_by":12,"title":"Figure 12","display":"","copyAsset":false,"role":"figure","size":179612,"visible":true,"origin":"","legend":"\u003cp\u003eComparison of developmental traits between \u003cem\u003eAcaulospora delicata\u003c/em\u003e-association and non-assosiation wheat plants under different water levels. (A) Plant fresh weight; (B) Plant dry weight. Yunhao-618: drought-resistant cultivar; Xinong-1376: drought non-resistant cultivar. Different letters on the bars indicate significant differences among treatments at the \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05 level, as determined by ANOVA followed by Tukey test. WDS: water-deficit stressed; WW: well-watered. Error bars represent the mean ± SE.\u003c/p\u003e","description":"","filename":"image12.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8715853/v1/478733cff697c23f8c046bd0.jpeg"},{"id":101754867,"identity":"0a294754-a45d-4996-aa08-cd13786bba16","added_by":"auto","created_at":"2026-02-03 10:47:27","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":3283258,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8715853/v1/2e7db88f-2f93-4388-a1fd-d7061c5f2a57.pdf"}],"financialInterests":"The authors declare no competing interests.","formattedTitle":"\u003cp\u003e\u003cstrong\u003eArbuscular mycorrhizal fungus \u003c/strong\u003e\u003cem\u003e\u003cstrong\u003eAcaulospora delicata \u003c/strong\u003e\u003c/em\u003e\u003cstrong\u003emodulates tri-trophic interactions between wheat cultivars and \u003c/strong\u003e\u003cem\u003e\u003cstrong\u003eSitobion avenae\u003c/strong\u003e\u003c/em\u003e\u003cstrong\u003e under drought stress\u003c/strong\u003e\u003c/p\u003e","fulltext":[{"header":"Introduction","content":"\u003cp\u003eClimate change is making droughts more frequent, persistent and severe, exacerbating droughts caused by insufficient rainfall or snowfall, leading to water scarcity, soil drought and crop damage (NASA \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; UNICEF \u003cspan citationid=\"CR118\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). These are the main causes of drought stress and the most critical abiotic stresses affecting global agricultural production such as wheat, with crop yields falling by 60% or more in hard-hit areas (Sareen et al. \u003cspan citationid=\"CR100\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Nyaupane et al. \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Wheat (\u003cem\u003eTriticum aestivum\u003c/em\u003e) is a major cereal crop that is important to the world's food supply and nutritional security for several hundred million people around the world (Rakszegi et al. \u003cspan citationid=\"CR95\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Endalew et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Wheat is highly susceptible to water scarcity, which can alter its physiological and metabolic processes, leading to growth damage, reduced biomass, and decreased grain quality (Filip et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Elnajar et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Wheat crops are also susceptible to various biotic stresses, including insect pests. Aphids, especially wheat aphids, are one of the most destructive pests of wheat, causing direct damage through feeding on the phloem and indirect damage through vector plant viruses such as barley yellow dwarf virus (Ding et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Leybourne et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) and excretes a sticky substance called honeydew, which is a food source for sooty mold and promotes their growth. Sooty mold is a black fungus that appears on the ears, rather than on other surfaces of wheat leaves and plants (Sabri et al. \u003cspan citationid=\"CR98\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Talabac \u003cspan citationid=\"CR112\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Lee \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), significantly reduced their photosynthetic capacity, thereby harming the health of plants. Aphid infestation can significantly reduce wheat productivity, with yield losses ranging from 10% to 70%, depending on the severity of the infection and the health status of the plant (Yahya et al. \u003cspan citationid=\"CR125\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eArbuscular mycorrhizal fungi (AMF) have established widespread attention for their potential to enhance plant tolerance to abiotic, biotic stresses (Ahammed et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) and forms a binding relationship with plant roots, extending hyphae into the soil and promoting greater water and nutrient absorption (Liu et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2018b\u003c/span\u003e). In return, the plants provide the fungi with carbohydrates produced via photosynthesis (Gunjal \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). In addition to their role in nutrient acquisition, AM fungi have been shown to enhance plant drought tolerance by increasing root hydraulic conductivity, enhancing osmotic regulation, and strengthening the synthesis of drought related hormones (Diagne et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Ye et al. \u003cspan citationid=\"CR130\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The role of AMF affect plant defense against herbivores by regulating primary and secondary metabolic pathways in plants (Amani et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Orine et al. \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). AMF, \u003cem\u003eAcaulospora delicata\u003c/em\u003e, is a group of fungi that inhabit soil and establish reciprocal relationships with the roots of most terrestrial plants, including wheat, to increase resistance to various abiotic and biotic stresses. For instance, wheat plants colonized by \u003cem\u003eA. delicata\u003c/em\u003e are better equipped to cope with drought conditions due to increased nutrient uptake, particularly Phosphorus, and improved water uptake and retention through an extensive hyphal network (da Trindade et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). The colonization may affect the interaction between plants and pests by altering the plants\u0026rsquo; nutritional status and defense mechanisms, thereby increasing or decreasing their susceptibility to herbivores such as \u003cem\u003eS. avenae\u003c/em\u003e (Koricheva et al. 2009). However, the effects of \u003cem\u003eA. delicata\u003c/em\u003e on wheat's interactions with \u003cem\u003eS. avenae\u003c/em\u003e under water-stress conditions have not been adequately studied.\u003c/p\u003e \u003cp\u003eIt is crucial to study the role of AMF, especially \u003cem\u003eA. delicata\u003c/em\u003e, in regulating wheat aphid interactions under drought conditions for several reasons. Firstly, it can provide insights into sustainable pest management strategies by utilizing natural plant fungal associations rather than relying solely on chemical insecticides. This sustainable approach is increasingly necessary due to growing concerns about pesticide resistance among pests and the negative environmental impacts associated with chemical use (Zou et al. \u003cspan citationid=\"CR135\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Secondly, understanding the comprehensive impact of drought resistant cultivars and AMF on aphid performance will provide information for cultivating more stress resistant wheat cultivars. These cultivars may utilize the benefits of AMF combination to enhance drought resistance and resistance to aphid infestation (Madouh et al. 2023). Thirdly, this study will contribute to a broader understanding of how global climate change will affect the dynamics of plant pests, and it is expected that global climate change will increase the frequency and severity of droughts. Investigating the interactions between AMF, wheat, and aphids under conditions of water scarcity will enable better predictions and mitigation strategies for the possible impacts of climate change on agricultural systems (Tang et al. \u003cspan citationid=\"CR113\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; George and Ray \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Finally, the findings from this study could have broader implications across other cereal crops and pest interactions. Understanding the mechanisms that govern AMF-plant-insect interactions may reveal similarities that can be applied to various species, facilitating enhanced agricultural practices (Pons et al. \u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Wahab et al. \u003cspan citationid=\"CR121\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The interaction between wheat cultivars, pests, and water stress is complex (Khoso et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2025a\u003c/span\u003e). Drought resistant wheat cultivars may enhance their defense against pests such as aphids, possibly due to physiological and morphological adaptations to water stress (Li et al. \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). However, it remains unclear how biotic interactions, particularly with aphids, shift when water stress is involved. The role of associative relationships, such as the association between wheat roots and AMF, under these stressful conditions has been limitedly explored (Rahman et al. \u003cspan citationid=\"CR94\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). It is known that \u003cem\u003eA. delicata\u003c/em\u003e enhances plant resistance to abiotic stress by increasing nutrient and water absorption (Carre\u0026oacute;n-Abud and Gavito \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Zhang et al. \u003cspan citationid=\"CR134\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), but its role in regulating wheat aphid interactions under drought stress is not fully understood.\u003c/p\u003e \u003cp\u003eThis study aims to explore the effects of wheat cultivars and \u003cem\u003eA. delicata\u003c/em\u003e on the performance of \u003cem\u003eS. avenae\u003c/em\u003e under water-deficit stress, with a focus on several key research objectives: 1) Quantify root length colonization of \u003cem\u003eA. delicata\u003c/em\u003e in different wheat cultivars under well-watered and water-deficit conditions. 2) Assess the effects of \u003cem\u003eA. delicata\u003c/em\u003e symbiosis on key life history traits of \u003cem\u003eS. avenae\u003c/em\u003e, including developmental duration, longevity, fecundity, and adult weight, across water regimes and cultivars. 3) Measure aphid honeydew production and evaluate changes in aphid fresh and dry mass associated with \u003cem\u003eA. delicata\u003c/em\u003e under drought stress. 4) Determine aphid host choice preference between drought-resistant and drought-susceptible wheat cultivars with and without \u003cem\u003eA. delicata\u003c/em\u003e symbiosis. 5) Evaluate the impact of \u003cem\u003eA. delicata\u003c/em\u003e on wheat root and shoot biomass under aphid infestation in both well-watered and water-deficit environments.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003e2.1 Aphids and plants\u003c/h2\u003e \u003cp\u003eThe anholocyclic \u003cem\u003eS. avenae\u003c/em\u003e clones were collected near the campus of Northwest A\u0026amp;F University. These clones were then reared on the Aikang-58 variety of wheat under environmental control condition (16L: 8D; 22\u0026thinsp;\u0026plusmn;\u0026thinsp;2\u0026deg;C). The seeds of wheat cultivars Yunhao-618 and Xinong-1376 were used in this study. Seeds were sown in plastic cups, which contained a mixture of turfy soil, vermiculite, and perlite in a 4:3:1 volumetric ratio. This mixture offers several benefits for plant growth (Supreme \u003cspan citationid=\"CR111\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The plastic cups were covered with transparent plastic tubes that were 7 cm in diameter and 20 cm in height. These tubes had a net on top with a 60-mesh size for ventilation purposes. This setup ensured optimal growth conditions by balancing moisture retention, drainage, and aeration, coupled with adequate light exposure and ventilation.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003e\u003cb\u003e2.2 Management of water conditions\u003c/b\u003e\u003c/h2\u003e \u003cp\u003eIn this study, we managed two water levels well-watered and water-deficit stress, as described previously in Dai et al. (\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Concisely, wheat seedlings were treated with 21 ml and 7 ml of water, respectively, every 4\u0026ndash;5 days. The seedlings were grown in 250 g (air-dry weight) of a growing medium consisting of a mixture of autoclaved quartz sand and soil inoculated with \u003cem\u003eA. delicata\u003c/em\u003e. For the non-association control, experimental cups did not contain the AMF \u003cem\u003eA. delicata\u003c/em\u003e inoculum with the growing media (a mixture of quartz-sand and soil). Required water conditions were maintained by measuring soil water potential and weighing the plants with the growing medium weekly by following the protocols described in Liu et al. (\u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2018a\u003c/span\u003e). Small stainless-steel pans were placed at the base of all cups to deliver an appropriate volume of water as needed.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003e\u003cb\u003e2.3 Propagation of the arbuscular mycorrhizal fungus\u003c/b\u003e\u003c/h2\u003e \u003cp\u003eThe AMF \u003cem\u003eAcaulospora delicata\u003c/em\u003e (BGCBJ02B) was obtained from the College of Natural Resources and Environmental, Northwest A\u0026amp;F University, China. For propagation, local maize (\u003cem\u003eZea mays\u003c/em\u003e) was used as the host plant. The selected propagation medium consisted of 25% soil and 75% quartz sand, ensuring an optimal environment for AMF development. To ensure a contamination-free environment, the surface of 25 maize seeds was sterilized using a combination of sodium hypochlorite and ethanol. The given method for sterilizing maize seeds follows to established protocols, involving a 70% ethanol soak for 2\u0026ndash;3 minutes followed by a 5% sodium hypochlorite soak for 5 minutes, followed by thorough rinsing with sterile water, to prevent contamination in subsequent stages of fungal inoculation and plant growth. This multi-step sterilization process helps to eliminate any potential pathogens while maintaining seed viability. Post-sterilization, the seeds were placed on a moist filter paper within a Petri dish (150 mm in diameter). The moisture levels in the Petri dish were meticulously maintained until germination occurred, ensuring that the seeds had adequate water for successful sprouting. An 8-liter plastic container (dimensions: 27\u0026times;20\u0026times;17 cm) was used to house the pre-germinated maize seeds. The container was filled with a growth medium composed of 8 kilograms of pasteurized loam and sand in a 1:3 volumetric ratios. Pasteurization of the medium is essential to eliminate any unwanted organisms that could interfere with AMF propagation. On top of the growth medium, 500 grams of \u003cem\u003eA. delicata\u003c/em\u003e fungal inoculum containing spores and root fragments was added. This inoculum was thoroughly mixed with the top layer of the medium to ensure even distribution. To complete the setup, an additional 500 grams of the sterilized medium was added on top, covering the inoculum and providing a protective layer. The pre-germinated maize seeds were carefully placed on the prepared medium in the 8-liter plastic containers. The mixture of pasteurized loam and sand, combined with the \u003cem\u003eA. delicata\u003c/em\u003e inoculum, provides a conducive environment for both the host plant and the fungus. Additionally, the layering technique ensures that the AMF inoculum is well-integrated within the medium to promote symbiosis between the maize roots and the mycorrhizal fungus (Parnell et al., \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The containers were placed in a climate chamber that provided controlled environmental conditions for four months under the following conditions: 16:8-hour light/dark cycle was employed to simulate natural light conditions, the temperature in the climate chamber was maintained at a range of 22\u0026thinsp;\u0026minus;\u0026thinsp;20℃, and the relative humidity was kept constant at 65% (Telli et al. \u003cspan citationid=\"CR114\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The plants were watered with the Hoagland nutrient solution described by Hoagland and Snyder (\u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e1934\u003c/span\u003e), once a week. The colonization of maize roots by the arbuscular mycorrhizal fungus \u003cem\u003eA. delicata\u003c/em\u003e was evaluated. The presence of mycorrhizal spores was confirmed by using the sieving method as described by Gerdemann and Nicholson (\u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e1963\u003c/span\u003e). This method involves filtering the soil sample through filter paper, followed by transferring the filtered material to a Petri dish where the number of spores is counted. This technique is essential for accurately assessing the spore presence, thereby confirming mycorrhizal colonization levels in the root samples.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003e2.4 Arbuscular mycorrhizal inoculation of wheat plants\u003c/h2\u003e \u003cp\u003eDrought-resistant (Yunhao-618) and drought non-resistant (Xinong-1376) cultivars of wheat were used in this experiment. The surface seed sterilization process was carried out using the method described above. After surface sterilization, both cultivars were placed on clean and moist filter paper in the petri dish (150 mm in diameter) under controlled environmental conditions of 22\u0026thinsp;\u0026plusmn;\u0026thinsp;2℃ (Zhang et al. \u003cspan citationid=\"CR133\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The filter paper was moistened with distilled water, and the petri plates were sealed with parafilm to maintain moisture levels. Upon successful germination, pre-germinated seeds were transferred to transparent plastic cups (7 cm in diameter). One pre-germinated seed was sown in each cup, ensuring careful handling to avoid damage to the emerging radicle (Rachel \u003cspan citationid=\"CR93\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Each cup contained approximately 100 g mixture of autoclaved quartz sand and soil. The quartz sand was sterilized at 130℃ for 30 minutes using an autoclave and allowed to cool down for 12 hours before the experiment (Scott \u003cspan citationid=\"CR103\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). This ensures the elimination of microbial contaminants that could interfere with the experiment. Next, a layer of 25 g of \u003cem\u003eA. delicata\u003c/em\u003e inoculum was added to the soil in each cup. A pre-germinated seed was then inserted on top of this layer of inoculum, followed by covering the seed with approximately 25 g of soil (Hibilik et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). This setup was designed to promote the establishment of an associative relationship between the wheat plant roots and the mycorrhizal fungus. At the end of the experiment, the roots were carefully extracted from the soil (Khoso et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2025b\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003e\u003cb\u003e2.5 Arbuscular mycorrhizal fungus root cleaning and assessment of fungal colonization\u003c/b\u003e\u003c/h2\u003e \u003cp\u003eThe roots of \u003cem\u003eA. delicata\u003c/em\u003e-inoculated wheat were initially removed from cups, thoroughly washed with regular tap water, and cut into segments 0.5-1 cm in length. These segments were then placed in 10 ml centrifuge tubes containing a weak formalin-acetic acid-alcohol (FAA) solution, composed of 50 ml formalin, 50 ml glacial acetic acid, and 900 ml ethyl alcohol, and stored at room temperature until further processing. After FAA treatment, the root segments underwent a soaking process in a 10% potassium hydroxide (KOH) solution (m/v) for 1 hour at 90\u0026deg;C to facilitate clearance and softening. Following the KOH treatment, the segments were rinsed with tap water before being subjected to an additional bleaching step. This involved immersion in a 10% alkaline solution at 20\u0026deg;C for 30 minutes, ensuring that any residual alkaline was thoroughly removed through multiple rinses. Subsequent to the bleaching process, the root segments were acidified by soaking them in a 1% hydrochloric acid (v/v) solution for a brief period of 3\u0026ndash;5 minutes. This acidification prepared the roots for the staining procedure. The segments were then stained using a 0.01% fuchsine acid dye at 90\u0026deg;C for 10 minutes. Following the staining, the roots were decolorized using a lactophenol staining solution, allowing for satisfactory visibility of the fungal structures within the roots. After cleaning and staining, the root segments were prepared for the detection and assessment of \u003cem\u003eA. delicata\u003c/em\u003e colonization following the methods described by Khoso et al. (\u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2025b\u003c/span\u003e). The processed root samples were taken from the centrifuge tubes and placed in glass petri dishes for de-staining and observation. Ten 1 cm-long root segments from both \u003cem\u003eA. delicata\u003c/em\u003e symbiosis treatments and non-symbiosis controls were randomly selected and mounted on biological microscope slides with coverslips for examination. The prepared slides were examined under an Olympus microscope (Olympus Model: U-LH100-3 Olympus Corporation, Tokyo 163\u0026ndash;0914, Japan) equipped with imaging software (cellSens Ver.1.7.1). The total root length colonization (TRLC) was estimated according to the methodology provided by Khoso et al. (\u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2025b\u003c/span\u003e). The intensity of \u003cem\u003eA. delicata\u003c/em\u003e colonization on individual root segments was evaluated using a rating scale from 0\u0026ndash;5 (Kormanik and McGraw \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e1982\u003c/span\u003e), with calculations performed using the Mycocalc software \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www2.dijon.inra.fr/mychintec/Mycocalc-prg/download.html\u003c/span\u003e\u003cspan address=\"https://www2.dijon.inra.fr/mychintec/Mycocalc-prg/download.html\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) according to Trouvelot et al. (\u003cspan citationid=\"CR117\" class=\"CitationRef\"\u003e1986\u003c/span\u003e), enabling precise quantification of colonization parameters.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003e2.6 Aphid performance bioassays\u003c/h2\u003e \u003cp\u003eHealthy \u003cem\u003eS. avenae\u003c/em\u003e adults were first transferred onto new wheat seedlings. On the next day, single newly emerged neonate nymphs (\u0026lt;\u0026thinsp;24 h old) were transferred to wheat seedlings of each cultivar in plastic cups (7 cm in diameter, one nymph per cup) using a camel hair brush. These plastic cups were well covered with transparent plastic tubes (30 cm in height, 6 cm in diameter), which had a net on top (60 mesh) for ventilation. Stainless steel pans were placed at the base of all cups to provide a suitable amount of water when needed. The fecundity, developmental time, and adult body weight of \u003cem\u003eS. avenae\u003c/em\u003e fed wheat were determined using the following steps. After pre-germination of the seeds, wheat seedlings were cultured in plastic cups filled with a medium consisting of a mixture of 25% soil and 75% quartz sand. This medium was sterilized at 130\u0026deg;C for 30 minutes and included 25 g of \u003cem\u003eA. delicata\u003c/em\u003e inoculum (Pokorny \u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Newly born nymphs of \u003cem\u003eS. avenae\u003c/em\u003e were placed on 7-day old wheat plants (1st wheat leaf) in plastic cups. Teneral adults were weighed on a micro-balance (METTLER-TOLEDO, XS3DU, Greifensee, Switzerland) and then transferred back to the test plants. These adults were left undisturbed and monitored daily until all test aphid individuals died. Newly born aphids were removed every day to avoid overcrowding. To ensure reliability, 40 replicates were observed for each wheat cultivar grown under both water-deficit-stressed and well-watered conditions.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec9\" class=\"Section2\"\u003e \u003ch2\u003e2.7 Honeydew excretion assay\u003c/h2\u003e \u003cp\u003eIn this assay, we compared the mass of honeydew excreted by the aphids, \u003cem\u003eS. avenae\u003c/em\u003e, when fed on drought-resistant Yunhao-618 and drought-non-resistant Xinong-1376, under well-watered and water deficit-stressed conditions. Wheat seeds were planted as above mentioned. One apterous \u003cem\u003eS. avenae\u003c/em\u003e adult was restricted to the first leaf of each wheat plant using a self-made clip cage (Kou et al., \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). The self-made clip cages were essential for confining aphids and ensuring accurate observation of honeydew production. After 24 hours, all young nymphs produced by the adult aphid were discarded to prevent overcrowding and ensure the accuracy of data. Following the initial setup period, a small piece of dried Jinjiayou aluminum foil (3 cm in diameter) Model: 613 (manufactured by Dongguan Juyou Paper \u0026amp; Plastic Products Co., Ltd, Guangdong China) was placed around the aphid-settled leaf inside each clip cage for an additional 24 hours. This setup ensured that honeydew excreted by the aphid would fall onto the aluminum foil for collection. The aluminum foil paper was dried to a constant weight in a drying oven at 60℃ for 30 minutes both before and after honeydew collection. This step was crucial to ensure that the weight of the honeydew could be accurately measured. The collected honeydew was then weighed using a microbalance, specifically the METTLER-TOLEDO XS3DU, which has a readability of 1 microgram. This precise measurement enabled an accurate comparison of honeydew excretion between the two wheat cultivars. A total of 40 replications were carried out for each wheat cultivar under both well-watered and water deficit-stressed conditions. Replication is a fundamental aspect of experimental design, ensuring the reliability and validity of the results.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec10\" class=\"Section2\"\u003e \u003ch2\u003e2.8 Aphid host choice\u003c/h2\u003e \u003cp\u003eThe experimental structure consisted of two 30 cm\u0026sup3; cages, connected by a 60 cm \u0026times; 15 cm transparent plastic tunnel. Each trial involved the use of one plant cultivar at a time, with 10 \u003cem\u003eA. delicate\u003c/em\u003e-inoculated plants placed in one cage and 10 non-symbiosis control plants placed in the other cage. The experimental design incorporated regular switching of the positions of the two cages to avoid positional effects, ensuring consistent environmental conditions for both treatment groups throughout the experiment (Guo et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Approximately fifty alate aphids were released in the center of the transparent plastic tunnel (4 cm in diameter) for each trial. This strategic positioning in the middle of the tunnel allowed the aphids to distribute themselves freely between the two cages. The numbers of aphids that settled within each cage were recorded at specific intervals: 2, 4, 8, 24, 48, and 72 hours post-release. All aphid individuals on a plant within a cage were considered settlements. The experimental design focused on recording the number of aphids settled on plants with AMF symbiosis (\u003cem\u003eA. delicata\u003c/em\u003e\u0026ndash;inoculated) and those without (non-symbiosis control). The consistent timing intervals allowed for detailed observation of aphid preferences over time, contributing to the understanding of how AMF symbiosis influences aphid host choice. Detailed comparative data was collected for each time point to analyze trends and patterns in aphid behavior. To ensure the reliability and robustness of the results, the experiment was conducted with a total of 20 replicates for each treatment. This high level of replication provided a robust dataset, allowing for statistical analysis to deduce significant differences and patterns in aphid settlement behavior between the two wheat cultivars.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003e2.9 Aphid desiccation assay\u003c/h2\u003e \u003cp\u003eUsing the above-mentioned method, aphid individuals of \u003cem\u003eS. avenae\u003c/em\u003e were raised on drought-resistant (Yunhao-618) and drought-non-resistant (Xinong-1376) wheat cultivars under well-watered and water deficit stressed levels for the bioassays of desiccation traits. A transparent desiccator glass pot, measuring 12\" \u0026times;16\", was used to maintain \u003cem\u003eS. avenae\u003c/em\u003e individuals under desiccation stress. The desiccator is an airtight chamber designed to regulate humidity levels effectively, making it ideal for experiments requiring controlled moisture conditions. The pot was carefully covered with a lid to ensure an airtight seal, which is essential to prevent atmospheric moisture from entering the chamber. To maintain the desired relative humidity within the desiccator, a layer of silica gel was used. Non-toxic and suitable silica gel, no risk of chemical interaction (purchased from Shandong Rushan Taihe Co., Ltd.), to help maintain the desired relative humidity. The desiccator pot was equipped with a digital hygrothermograph recorder (Anymeter, TH20, Guangzhou, Guangdong, China), to provide real-time monitoring of temperature and humidity. For each trial, a total of ten aphid individuals were put into a 10 ml centrifuge tube. The centrifuge tube was modified to create a hole and had a net (60 mesh) to cover the hole for ventilation. This setup ensured adequate airflow and prevented the aphids from escaping. Test aphid individuals were maintained under a specific environmental condition, which in this case was 10% relative humidity (RH), for about 12 hours. During this period, the survival rates of the aphids were meticulously recorded to assess the impact of low humidity on their viability (Yang et al. \u003cspan citationid=\"CR129\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). We weighed the test \u003cem\u003eS. avenae\u003c/em\u003e adults on a microbalance (METTLER-TOLEDO, XS3DU, Greifensee, Switzerland) to determine their fresh body mass (Mf). After they were subjected to the desiccation stress treatment for 12 h in a desiccator, test aphid individuals were reweighed to obtain the body mass after the desiccation exposure of 12 h (M\u003csub\u003e12\u003c/sub\u003e h). Thus, the rate of water loss per hour in this study refers to the mean water loss during the initial 12 hours of desiccation stress. After they were dead, test aphid individuals were dried at 60℃ for 60 h and then weighed again to obtain the dry mass (Md). We estimated the water loss rates using the exponential model M\u003csub\u003e12\u003c/sub\u003eh\u0026thinsp;=\u0026thinsp;Mf e-\u003csup\u003ekt\u003c/sup\u003e, and calculated the total water loss rates as kt = -ln (M\u003csub\u003e12\u003c/sub\u003eh/M\u003csub\u003ef\u003c/sub\u003e) (Wharton \u003cspan citationid=\"CR124\" class=\"CitationRef\"\u003e1985\u003c/span\u003e). Following Gibbs et al. (\u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e1997\u003c/span\u003e), absolute water contents were calculated as M\u003csub\u003ef\u003c/sub\u003e- M\u003csub\u003ed\u003c/sub\u003e and relative water contents as (M\u003csub\u003ef-\u003c/sub\u003eM\u003csub\u003ed)/\u003c/sub\u003eM\u003csub\u003ef\u003c/sub\u003e\u0026times;100).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003e2.10 Plant developmental traits\u003c/h2\u003e \u003cp\u003eAM fungus inoculation occurred at the time of sowing seeds. First, cups of 4 cm diameter were filled with approximately 175 g of sand. Then 50 g of inoculum was mixed with the top layer of the soil, and a pre-germinated seed was added on top of the inoculum-mixed soil, which was covered on top with approximately 25 g medium (sand and soil). The control (i.e., non-symbiosis) plants were treated identically but used inoculum that had been autoclaved and allowed to dry for 24 hours before sowing. All the treated wheat plants were kept in a controlled environmental room (21\u0026thinsp;\u0026plusmn;\u0026thinsp;2℃, relative humidity of 65\u0026thinsp;\u0026plusmn;\u0026thinsp;5%), and the cups were placed in a randomized block design with a total of 40 replications for each treatment. Plant heights were measured, and numbers of leaves were counted weekly. At the end of the experiment, all plant samples were taken from the cups. Before drying the plant material, fresh and dry weights of whole plants were obtained by using an electronic analytical balance (Sartorius TE20101-L). The shoots, stems and leaves were oven-dried at 75℃ for 60h. Ten replicates were performed for each treatment.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003e3.11 Statistical analyses\u003c/h2\u003e \u003cp\u003eA randomized block (i.e., batch) design was used for the experiments. The bioassays were conducted in two batches (five replicates per treatment in each batch), with test aphid individuals randomly assigned to the treatments (i.e., symbiosis, water level and cultivar) within each batch. Ten replicates were conducted for each treatment. The data on mycorrhizal root colonization were compared with two-way ANOVA, and the effects of independent factors (i.e., cultivar and water level) were analyzed, as well as the interaction between cultivar and water level. Similarly, the total amino acid contents of wheat plants were examined with two-way ANOVA. We used three-way ANOVA to analyze the data on the developmental durations of nymphal instars, adult weight, fecundity, plant development parameters (i.e., fresh and dry weight), honeydew weight, and aphid desiccation as response variables, and the effects of independent factors, including symbiosis, water level and cultivar, were analyzed, as well as the interactions among the three factors. Mean separations were done with Tukey HSD (honestly significant differences) tests after significant ANOVA. We used the independent sample student\u0026rsquo;s \u003cem\u003et\u003c/em\u003e-test to identify significant differences between the association treatment and the non-association control (\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05) for settlement of \u003cem\u003eS. avenae\u003c/em\u003e over the course of 72 hours. Data were analyzed by using Statistix 8.1, and all figures were generated by using the OriginPro 2024 software.\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003e3.1 AMF \u003cem\u003eA. delicata\u003c/em\u003e percentage root length colonization (%RLC)\u003c/h2\u003e \u003cp\u003eAMF \u003cem\u003eA. delicata\u003c/em\u003e RLC was observed in drought resistant (Yunhan-618) and drought resistant (Xinnong-1376) wheat cultivars under different water conditions. For RLC, it was found that water level has a significant impact (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e; \u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;334.47; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1.72;\u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.001). Under water deficit-stress, high percentage of AMF \u003cem\u003eA. delicata\u003c/em\u003e spore population was recorded in Xinong-1376 and Yunhao-618 cultivars with 67% and 62%, respectively. While control plants did not show any associations with spore production. Xinong-1376 and Yunhao-618 plants may form beneficial associations with AMF, which significantly enhances their drought tolerance. The colonization \u003cem\u003eA. delicata\u003c/em\u003e was clearly shown in the fungus inoculated wheat plants (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eA), as compared to control plants exhibiting zero colonization (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eB).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cb\u003e3.2 Life history traits of\u003c/b\u003e \u003cb\u003eS. avenae\u003c/b\u003e \u003cb\u003eaphids under the influence of mycorrhizal fungus\u003c/b\u003e \u003cb\u003eA. delicata\u003c/b\u003e \u003cb\u003eassociations\u003c/b\u003e\u003c/p\u003e \u003cdiv id=\"Sec16\" class=\"Section3\"\u003e \u003ch2\u003e3.2.1 Developmental duration\u003c/h2\u003e \u003cp\u003eDevelopmental duration (DD) of \u003cem\u003eS. avenae\u003c/em\u003e aphid, the research shows significant effects of various factors on its DD, including cultivars, water levels, and mycorrhizal fungus \u003cem\u003eA. delıcata\u003c/em\u003e associations. The effects of association on first instar nymphs that the DD for the first instar nymphs of \u003cem\u003eS. avenae\u003c/em\u003e prolonged with mycorrhizal non-associations on Xinong-1376 plants under water-deficit conditions. Conversely, these nymphs exhibit shorter under well-watered conditions on the Yunhao-618 cultivar. For DDs of first instar nymphs, there were no significant interactions between test factors identified. The significant effects of these conditions (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eA; cultivars: \u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;7.14; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05; associations: \u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;12.21; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001; and water levels: \u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;18.63; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001) for aphid development in varying moisture environments. The DD of second instar \u003cem\u003eS. avenae\u003c/em\u003e nymphs was significantly affected by AMF \u003cem\u003eA. delicata\u003c/em\u003e association. Significant interactions were found between mycorrhizal association and cultivars (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eB; \u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;14.58; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001); and their interactions between test factors (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;6.48; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.05) identified. The association with AMF \u003cem\u003eA. delicata\u003c/em\u003e significantly affected the development duration of second instar nymphs (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;8.82; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05). For the DD third instar nymphs of \u003cem\u003eS. avenae\u003c/em\u003e, there were significant effects of different cultivars (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eC; \u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;9.42; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05). For DD of fourth instar nymphs, there were no significant interactions between test factors identified. The association treatments showed longer DD when associated with mycorrhizal fungi on Yunhao-618 plants under well-watered conditions. In contrast, the DD on Xinong-1376 under water deficit stressed conditions was notably shorter (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eD). Significant interactions between cultivar and association were identified for the total developmental time of \u003cem\u003eS. avenae\u003c/em\u003e nymphs (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e; \u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;5.83; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05), and cultivar and water levels (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;4.00; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05). The total DD for \u003cem\u003eS. avenae\u003c/em\u003e under well-watered conditions was longer (9.5 days) on mycorrhizal non-associated Xinong-1376 plants compared to a shorter DD (7 days) under water deficit-stressed conditions.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec17\" class=\"Section3\"\u003e \u003ch2\u003e3.2.2 Aphid longevity\u003c/h2\u003e \u003cp\u003eThe longevity of \u003cem\u003eS. avenae\u003c/em\u003e on plants without \u003cem\u003eA. delicata\u003c/em\u003e association was shorter compared to those with the association across both cultivars and water treatments. It was significant interactions between association and cultivar (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e; \u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;8.96; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05), and water level (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;85.52; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), but no significant interactions were found for this aphid. For example, under water-deficit conditions, the longevity of \u003cem\u003eS. avenae\u003c/em\u003e on Xinong-1376 was significantly shorter for the \u003cem\u003eA. delicata\u003c/em\u003e non-association control (mean\u0026thinsp;=\u0026thinsp;13.8 d) compared to the association treatment (mean\u0026thinsp;=\u0026thinsp;24.9 d). Similarly, under well-watered conditions, \u003cem\u003eS. avenae\u003c/em\u003e on Yunhao-618 had shorter longevity in the non-association control (mean\u0026thinsp;=\u0026thinsp;14.7 d) compared to the association treatment (under well-watered: mean\u0026thinsp;=\u0026thinsp;21.8 d; and under water deficit-stressed: mean\u0026thinsp;=\u0026thinsp;21.6 d).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec18\" class=\"Section3\"\u003e \u003ch2\u003e3.2.3 10-d fecundity\u003c/h2\u003e \u003cp\u003eThe presence of AMF \u003cem\u003eA. delicata\u003c/em\u003e appears to beneficially affect \u003cem\u003eS. avenae\u003c/em\u003e fecundity (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e; \u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;10.73, \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001). Water conditions also play a crucial role in determining aphid fecundity (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;151.37, \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001). However, the association with \u003cem\u003eA. delicata\u003c/em\u003e seems to mitigate some of these negative effects, as indicated by the higher mean fecundity under water-deficit stress in plants associated with \u003cem\u003eA. delicata\u003c/em\u003e (mean\u0026thinsp;=\u0026thinsp;22.4, SE\u0026thinsp;=\u0026thinsp;1.00) compared to non-associated plants (mean\u0026thinsp;=\u0026thinsp;10.2, SE\u0026thinsp;=\u0026thinsp;0.85). The cultivar Yunhao-618 showed higher fecundities in association with \u003cem\u003eA. delicata\u003c/em\u003e compared to Xinong-1376. This indicates that genetic differences between cultivars can influence the effectiveness of the \u003cem\u003eA. delicata\u003c/em\u003e association in enhancing \u003cem\u003eS. avenae\u003c/em\u003e fecundity (interaction between cultivar and association: \u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;14.94; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec19\" class=\"Section3\"\u003e \u003ch2\u003e3.2.4 Adult weight\u003c/h2\u003e \u003cp\u003eIn the context of AMF \u003cem\u003eA. delicata\u003c/em\u003e association treatments, \u003cem\u003eS. avenae\u003c/em\u003e exhibited higher adult weights on both Yunhao-618 and Xinong-1376 cultivars under well-watered conditions. There were significant differences due to the association (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003e; \u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;27.23; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001) and water level (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;85.44; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 22; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001) factors. When comparing the performance of aphids on different cultivars, Xinnong-1376 showed higher adult body weight under water stress conditions, Specifically, the average adult weight of \u003cem\u003eS. avenae\u003c/em\u003e on Yunhao-618 under well-watered conditions was significantly higher in the association treatment (mean\u0026thinsp;=\u0026thinsp;590 \u0026micro;g, SE\u0026thinsp;=\u0026thinsp;8.59) than in the non-association control (mean\u0026thinsp;=\u0026thinsp;362.6 \u0026micro;g, SE\u0026thinsp;=\u0026thinsp;8.93). Similarly, under water-deficit stress, the teneral adult body weight on Xinong-1376 was higher in the association treatment (mean\u0026thinsp;=\u0026thinsp;665.40 \u0026micro;g, SE\u0026thinsp;=\u0026thinsp;11.88) than in the non-association control (mean\u0026thinsp;=\u0026thinsp;417.00 \u0026micro;g, SE\u0026thinsp;=\u0026thinsp;18.9). The association treatments led to enhanced adult weights under both well-watered and water deficit-stressed conditions, which highlights the beneficial role of symbiotic relationships in mitigating the adverse effects of water stress on aphid performance.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003c/div\u003e \u003cdiv id=\"Sec20\" class=\"Section2\"\u003e \u003ch2\u003e3.3 Aphid honeydew production\u003c/h2\u003e \u003cp\u003eFor honeydew production of this aphid, cultivars (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003e; \u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;14.89; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1,72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), water level (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;46.11; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1,72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), and for association (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;22.05; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1,72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001) showed significant effects, but the effects for the interaction between cultivars and water level were not significant (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1.87; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1,72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;0.05), or for cultivars and association (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.05; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1,72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;0.05). The amount of honeydew production by \u003cem\u003eS. avenae\u003c/em\u003e under well-watered conditions was significantly higher than under water deficit-stressed conditions on all test wheat cultivars. With \u003cem\u003eA. delicate\u003c/em\u003e association under both water levels, this aphid produced higher amounts of honeydew (mean\u0026thinsp;=\u0026thinsp;4.27 \u0026micro;g, SE\u0026thinsp;=\u0026thinsp;0.11) on drought non-resistant Xinong-1376 than on drought-resistant cultivars. There was lowest amounts of honeydew (mean\u0026thinsp;=\u0026thinsp;0.97 \u0026micro;g, SE\u0026thinsp;=\u0026thinsp;0.14) on drought resistant Yunhao-618 under water deficit-stressed conditions.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec21\" class=\"Section2\"\u003e \u003ch2\u003e3.4 Aphid desiccation assays\u003c/h2\u003e \u003cdiv id=\"Sec22\" class=\"Section3\"\u003e \u003ch2\u003e3.4.1 Fresh mass\u003c/h2\u003e \u003cp\u003eIn terms of fresh body mass, we found significant effects for cultivars (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e9\u003c/span\u003eA; \u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;25.49; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), water level (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;20.93; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), and interactions between cultivars and association (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;6.78; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05). The fresh body masses of \u003cem\u003eS. avneae\u003c/em\u003e on the drought resistant Yunhao-618 tended to be higher with \u003cem\u003eA. delicate\u003c/em\u003e association than those on Xinong-1376 test drought non-resistant cultivars. Under water-deficit stress, high fresh body masses (mean\u0026thinsp;=\u0026thinsp;375.77 \u0026micro;g, SE\u0026thinsp;=\u0026thinsp;21.65) on Yunhao-618 and without \u003cem\u003eA. delicate\u003c/em\u003e association lower (mean\u0026thinsp;=\u0026thinsp;197.80 \u0026micro;g, SE\u0026thinsp;=\u0026thinsp;5.15) on Xinong-1376, but in terms of \u003cem\u003eA. delicate\u003c/em\u003e association alone, this pattern was found opposite patterns on both cultivars.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec23\" class=\"Section3\"\u003e \u003ch2\u003e3.4.2 Dry mass\u003c/h2\u003e \u003cp\u003eFor the dry mass of this aphid, we identified significant effects for cultivars (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e9\u003c/span\u003eB; \u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;44.01; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), and water treatments (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;12.70; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), or for association (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;6.06; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05). On Yunhao-618, the dry body masses of this aphid tended to increase with increasing water-deficit stress, but this pattern was not found on other test cultivar (Xinong-1376). Under well-watered conditions, this aphid without \u003cem\u003eA. delicate\u003c/em\u003e association had higher dry body masses on Yunhao-618 than on Xinong-1376 cultivars.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec24\" class=\"Section3\"\u003e \u003ch2\u003e3.4.3 Water balance traits\u003c/h2\u003e \u003cp\u003eWe found effects on absolute water contents (AWC) of \u003cem\u003eS. avenae\u003c/em\u003e on cultivars (Fig.\u0026nbsp;\u003cspan refid=\"Fig10\" class=\"InternalRef\"\u003e10\u003c/span\u003eA; \u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;14.64; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), water level (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;19.28; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), and the interaction between cultivars and water level (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;6.60; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05), or interaction between cultivars and association (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;6.60; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05). The highest absolute water contents of this aphid were found with \u003cem\u003eA. delicate\u003c/em\u003e association on drought-resistant Yunhao-618 under water-deficit stressed conditions, but this aphid under water-deficit stressed had the lowest absolute water contents on Xinong-1376.\u003c/p\u003e \u003cp\u003eRelative water contents (RWC) of \u003cem\u003eS. avenae\u003c/em\u003e also showed significant variability under different conditions. There were significant effects identified for cultivars (Fig.\u0026nbsp;\u003cspan refid=\"Fig10\" class=\"InternalRef\"\u003e10\u003c/span\u003eB; \u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;24.04; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), association (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;11.23; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), and the interaction between cultivars and water level (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;26.54; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001). Additionally, significant interactions across all factors (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;4.52; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05) were observed. RWC of \u003cem\u003eS. avenae\u003c/em\u003e without \u003cem\u003eA. delicate\u003c/em\u003e association on drought resistant Yunhao-618 showed a decreasing trend with increasing well-water. Oppositely, this pattern was found on decreasing trend with increasing water deficit-stress on Xinong-1376 cultivar. This aphid had equal relative water contents on both cultivars with \u003cem\u003eA. delicate\u003c/em\u003e association under water deficit-stressed and well-watered conditions.\u003c/p\u003e \u003cp\u003eWater loss rates (WLR) in \u003cem\u003eS. avenae\u003c/em\u003e associated with \u003cem\u003eA. delicate\u003c/em\u003e showed significant interactions between cultivars and water level (Fig.\u0026nbsp;\u003cspan refid=\"Fig10\" class=\"InternalRef\"\u003e10\u003c/span\u003eC; \u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;11.64; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), as well as between cultivars and association (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;4.16; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05). Significant interactions between all factors were also noted (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;20.90; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001). Water loss rates of \u003cem\u003eS. avenae\u003c/em\u003e with \u003cem\u003eA. delicate\u003c/em\u003e association showed a decreasing trend with increasing water-deficit stress on Yunhao-618, and without association on Xinong-1376. Under well-watered, this aphid had the lowest water loss rates with \u003cem\u003eA. delicate\u003c/em\u003e association on Yunhao-618, and Xinong-1376 under water-deficit stressed.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003c/div\u003e \u003cdiv id=\"Sec25\" class=\"Section2\"\u003e \u003ch2\u003e3.5 Aphid host choice\u003c/h2\u003e \u003cp\u003eThe host choice performance of \u003cem\u003eS. avenae\u003c/em\u003e on different wheat cultivars under varying water conditions reveals complex dynamics influenced by the plant's drought resistance and association with AMF (\u003cem\u003eA. delicate\u003c/em\u003e). Under water-deficit stress, \u003cem\u003eS. avenae\u003c/em\u003e showed a significant preference for plants associated with \u003cem\u003eA. delicate\u003c/em\u003e over the non-association control on Yunhao-618 at 12 hours post-release (hpr) (Fig.\u0026nbsp;\u003cspan refid=\"Fig11\" class=\"InternalRef\"\u003e11\u003c/span\u003eA; \u003cem\u003et\u003c/em\u003e\u0026thinsp;=\u0026thinsp;7.40; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;18; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001) which was also observed under well-watered conditions (Fig.\u0026nbsp;\u003cspan refid=\"Fig11\" class=\"InternalRef\"\u003e11\u003c/span\u003eB; \u003cem\u003et\u003c/em\u003e\u0026thinsp;=\u0026thinsp;2.86; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;18; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05). A notable preference shift was seen in the Xinong-1376 cultivar under severe-water deficit at just 2 hpr (Fig.\u0026nbsp;\u003cspan refid=\"Fig11\" class=\"InternalRef\"\u003e11\u003c/span\u003eC; \u003cem\u003et\u003c/em\u003e\u0026thinsp;=\u0026thinsp;4.96; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;18; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001) and at 12 hpr under well-watered conditions (Fig.\u0026nbsp;\u003cspan refid=\"Fig11\" class=\"InternalRef\"\u003e11\u003c/span\u003eD; \u003cem\u003et\u003c/em\u003e\u0026thinsp;=\u0026thinsp;3.89, \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;18; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001). At 72 hpr, the aphid population with \u003cem\u003eA. delicate\u003c/em\u003e on Xinnong-1376 (under water-deficit stress mean: 19.20, SE\u0026thinsp;=\u0026thinsp;0.39; and under well-watered mean: 23.50, SE\u0026thinsp;=\u0026thinsp;0.23) was significantly higher than that of Yunhao-618 (under water-deficit stress mean: 5.50, SE\u0026thinsp;=\u0026thinsp;0.11; and under well-watered mean: 9.30, SE\u0026thinsp;=\u0026thinsp;0.26). At 72 hpr, \u003cem\u003eS. avenae\u003c/em\u003e showed with \u003cem\u003eA. delicate\u003c/em\u003e a strong preference for the drought non-resistant Xinong-1376 over the drought-resistant Yunhao-618 cultivar. It can be partly explained by the sustainability and nutrition provided by the respective plants under both water-deficit and well-watered conditions with \u003cem\u003eA. delicate\u003c/em\u003e. The mean aphid population was significantly higher on Xinong-1376.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec26\" class=\"Section2\"\u003e \u003ch2\u003e3.6 Plant developmental traits\u003c/h2\u003e \u003cp\u003eThe study found that wheat plants with AMF (\u003cem\u003eA. delicate\u003c/em\u003e) association treatments had significantly larger fresh weights compared to those without the associations (Fig.\u0026nbsp;\u003cspan refid=\"Fig12\" class=\"InternalRef\"\u003e12\u003c/span\u003eA; association: \u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;40.05; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001). Especially, under conditions of well-watered condition, the fresh plant weight of AMF combination is higher compared to water-deficit stressed conditions (water level: \u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;29.76; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1,72;\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001). The effect of plant dry weight on cultivars (Fig.\u0026nbsp;\u003cspan refid=\"Fig12\" class=\"InternalRef\"\u003e12\u003c/span\u003eB; \u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;5.56༛\u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1,72༛\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05), water level (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;11.80; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1,72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001) and association (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;5.20; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1.72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05) had a significant influence. In addition, the interaction between associations and cultivar (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;5.56; \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1, 72; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05) was significant.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003eThe high root length colonization (%RLC) by AMF \u003cem\u003eA. delicata\u003c/em\u003e observed in drought-resistant wheat cultivars Xinong-1376 (67%) and Yunhao-618 (62%) under water deficit stress underscores the important role of AMF in enhancing drought tolerance. This aligns with recent studies demonstrating that effective AMF colonization improves water and nutrient uptake, crucial for maintaining plant physiological functions during drought (Nopphakat et al. 2022; Akensous et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Similar findings show that AMF symbiosis increases root surface area and spore production, facilitating access to deeper soil moisture (Kang et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Li et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Furthermore, research highlights that AMF-mediated enhancements in antioxidant enzyme activity reduce oxidative stress under drought, contributing to improved plant resilience (Li et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Importantly, AMF colonization is cultivar-specific, influenced by root architecture and genetic compatibility, as observed in Xinong-1376 and Yunhao-618, suggesting targeted AMF inoculation could optimize drought adaptation (Duan et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). These synergistic insights affirm AMF \u003cem\u003eA. delicata\u003c/em\u003e as a promising bio-fertilizer to bolster wheat productivity in water-limited environments.\u003c/p\u003e \u003cp\u003eThe DD of \u003cem\u003eS. avenae\u003c/em\u003e nymphs was notably influenced by the combination of wheat cultivars, water availability, and association with AMF, particularly \u003cem\u003eA. delicata\u003c/em\u003e. Under water-deficit conditions, first instar nymphs showed prolonged DD on Xinong-1376 plants without AMF, indicating that drought stress and the absence of beneficial fungal symbiosis negatively impact aphid development (Babikova et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Liu et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2018b\u003c/span\u003e; Stallmann and Schweiger 2021). In contrast, under well-watered conditions, \u003cem\u003eYunhao-618\u003c/em\u003e plants with AMF supported faster development, suggesting that AMF may mitigate stress under optimal water availability (Thompson, \u003cspan citationid=\"CR115\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). The second instar nymphs exhibited longer DD on Xinong-1376 without AMF under well-watered conditions, while under drought, DD shortened\u0026mdash;suggesting stress may accelerate development in the absence of AMF. Similar findings by Kula et al. (\u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2005\u003c/span\u003e) and Cascone et al. (\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) show that AMF often reduce insect development time under optimal conditions by improving plant nutrition, but under stress, absence of AMF can lead to rapid aphid development. Cultivar selection also influenced aphid DD, though no significant interaction between cultivar and other factors was observed (Platkova et al. \u003cspan citationid=\"CR88\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). For third instar nymphs, \u003cem\u003eA. delicata\u003c/em\u003e shortened DD on Xinong-1376 under both water regimes, while under drought stress, Yunhao-618 without AMF caused significant DD prolongation. This suggests AMF can alleviate drought effects and enhance aphid development (Yang et al. \u003cspan citationid=\"CR128\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Liu et al. \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2018a\u003c/span\u003e). For fourth instars, longer DD was observed on Yunhao-618 with AMF under well-watered conditions, possibly due to AMF-induced plant defenses (Babikova et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Conversely, shorter DD under drought on \u003cem\u003eXinong-1376\u003c/em\u003e supports the idea that aphids may accelerate development to compensate for reduced plant quality (Hodge and Storer \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Gange et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Overall, aphid development is context-dependent, shaped by the complex interactions among plant genotype, AMF association, and water availability, consistent with findings that water stress and AMF play key roles in modulating aphid fitness and host plant quality (Beetge and Kr\u0026uuml;ger \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSignificant interactions between wheat cultivar and mycorrhizal association, as well as between cultivar and water levels, strongly influence the total developmental duration (DD) of \u003cem\u003eS. avenae\u003c/em\u003e nymphs. The longer DD (9.5 days) observed on mycorrhizal non-associated Xinong-1376 plants under well-watered conditions compared to the shorter DD (7 days) under water deficit stress aligns with recent studies showing that optimal water availability promotes aphid development by enhancing plant vigor and nutrient quality. Conversely, drought stress accelerates aphid development, possibly as a compensatory adaptation to reduced host quality. Similar findings by Liu et al. (2018) indicated that drought stress shortens aphid developmental periods, especially in the absence of beneficial fungal symbiosis. Moreover, AMF associations can moderate these effects by improving plant nutrition and stress tolerance, sometimes prolonging aphid development under favorable conditions (Babikova et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2014\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe observed increase in \u003cem\u003eS. avenae\u003c/em\u003e fecundity on wheat plants colonized by \u003cem\u003eA. delicata\u003c/em\u003e under water-deficit stress aligns with emerging research demonstrating that AMF can enhance host plant nutritional and physiological status, thereby benefiting phloem-feeding herbivores. Similar studies highlight AMF\u0026rsquo;s role in improving plant water and nutrient uptake under drought, which maintains or elevates phloem sap quality crucial for aphid reproduction (Gange \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Zeng \u003cspan citationid=\"CR132\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The cultivar-specific response, where Yunhao-618 shows heightened aphid fecundity under AMF symbiosis, underscores genetic variation in plant-mycorrhizal compatibility affecting herbivore performance (Khoso et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2025b\u003c/span\u003e). Contrastingly, research involving fungal endosymbionts in other aphid species found reduced fecundity and lifespan, suggesting that microbial associations can have positive or negative effects on herbivores depending on symbiont type and plant context (Meister et al. \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Johnson et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Additionally, drought typically reduces aphid fecundity and population growth (Kansman et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), but the buffering effect of \u003cem\u003eA. delicata\u003c/em\u003e suggests potential for leveraging AMF in integrated pest and drought management strategies to sustain crop resilience and manage aphid outbreaks effectively.\u003c/p\u003e \u003cp\u003eOur findings align with recent research highlighting the role of both wheat cultivar traits and AMF in shaping aphid performance under varying water conditions. Consistent with Liu et al. (\u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2018a\u003c/span\u003e), differences in adult \u003cem\u003eS. avenae\u003c/em\u003e weight across water treatments reflect the aphid\u0026rsquo;s adaptation potential, influenced by plant nutritional quality and stress resilience. AMF-driven enhancements in nutrient and water uptake improve host plant vigor, creating a more favorable environment for aphid growth, as seen in the higher adult weights on Xinong-1376 compared to Yunhao-618 under drought stress. This supports previous work showing drought-resilient cultivars maintain better aphid nutritional support (Pineda et al. \u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). However, severe water deficit generally reduces aphid fitness due to diminished plant resources, consistent with Stiling et al. (\u003cspan citationid=\"CR109\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). The positive influence of \u003cem\u003eA. delicata\u003c/em\u003e on aphid weight under both water regimes indicates AMF can buffer drought effects while simultaneously promoting aphid development, a dynamic critical for predicting pest outbreaks under climate change (Rizzo \u003cspan citationid=\"CR97\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). This interplay poses challenges for crop protection as increased aphid vigor may lead to more severe yield losses (Lenoir et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; CABI \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAphid honeydew production varies significantly with host plant cultivar, water availability, and AMF associations, consistent with findings from previous research. Well-watered plants generally support higher honeydew output due to increased nutrient availability and more efficient aphid feeding, aligning with Huberty and Denno (\u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2004\u003c/span\u003e) and van Rooijen (\u003cspan citationid=\"CR119\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Plant stress, such as drought, reduces phloem pressure and increases sap viscosity, hindering aphid feeding and lowering honeydew production, as observed by Kansman et al. (\u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Cultivar differences further influence these dynamics; drought-susceptible cultivars like Xinong-1376 provide more favorable feeding conditions for aphids regardless of water regime, leading to higher honeydew outputs. In contrast, resistant cultivars such as Yunhao-618 exhibit physical and chemical defenses that reduce aphid feeding efficiency and honeydew production under drought stress (Stallmann et al. 2022; Tous-Fandos et al. \u003cspan citationid=\"CR116\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). AMF association with \u003cem\u003eA. delicata\u003c/em\u003e enhances honeydew production, likely by improving phloem sap access or modulating plant defenses, supporting concepts from Styrsky and Eubanks (\u003cspan citationid=\"CR110\" class=\"CitationRef\"\u003e2007\u003c/span\u003e) and Milcu et al. (\u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). These insights emphasize the complex, genotype-environment-microbe interplay regulating aphid-plant interactions and pest dynamics under stress.\u003c/p\u003e \u003cp\u003eThe observed influence of \u003cem\u003eA. delicata\u003c/em\u003e symbiosis on aphid fresh and dry body mass under drought conditions echoes recent findings on the integral role of microbial associations in aphid physiology and adaptation. Similar to Csorba et al. (\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), symbiotic relationships with microbes such as \u003cem\u003eBuchnera aphidicola\u003c/em\u003e enhance aphid nutrient acquisition, critical under host plant stress. The cultivar-specific benefits, with Yunhao-618 facilitating higher aphid mass under drought, reflect differential plant physiological responses consistent with Pozo and Azc\u0026oacute;n-Aguilar (\u003cspan citationid=\"CR92\" class=\"CitationRef\"\u003e2007\u003c/span\u003e), who emphasize that drought-resistant cultivars better exploit AM fungi symbiosis to maintain plant health. Recent research also links increased aphid performance on drought-stressed resistant cultivars to altered phloem chemistry, such as elevated soluble sugars and nitrogen availability, supporting Huberty and Denno\u0026rsquo;s (\u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2004\u003c/span\u003e) and Kansman et al.\u0026rsquo;s (\u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) conclusions. Conversely, susceptible cultivars like Xinong-1376 experience compromised physiology under drought, reducing aphid support.\u003c/p\u003e \u003cp\u003eThe findings on the absolute water content (AWC), relative water content (RWC), and water loss rate (WLR) of \u003cem\u003eS. avenae\u003c/em\u003e reveal intricate interactions between aphids, host plant cultivars, water availability, and symbiotic association with \u003cem\u003eA. delicata\u003c/em\u003e. Echoing recent research on other herbivores such as \u003cem\u003eAphis nerii\u003c/em\u003e and leaf-cutting ants (Carvajal Acosta et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Gely et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), plant genotype and environmental stressors play pivotal roles in determining insect water status and physiological performance. The higher AWC and maintained RWC of \u003cem\u003eS. avenae\u003c/em\u003e on the drought-resistant cultivar Yunhao-618 under water deficit, especially when associated with AMF, align with studies demonstrating that mycorrhizal fungi enhance host plant water retention, nutrient uptake, and overall physiological resilience under drought (D\u0026iacute;az-Hern\u0026aacute;ndez et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Yan et al. \u003cspan citationid=\"CR126\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). These improvements indirectly benefit associated herbivores by stabilizing phloem sap quality and water availability. In contrast, drought-susceptible cultivars like Xinong-1376 show reduced aphid water content and higher water loss rates under stress, mirroring findings in other insect-plant systems where plant stress diminishes herbivore hydration and fitness (Benoit et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The mitigation of water loss in \u003cem\u003eS. avenae\u003c/em\u003e by AMF symbiosis suggests symbiotic fungi may contribute to improved insect desiccation tolerance, a trait critical under climate change-induced drought scenarios (Addo-Bediako et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). Moreover, the persistent aphid preference for AMF-associated plants across water regimes supports the hypothesis that mycorrhizal fungi improve plant nutritional quality and attractiveness independent of water status, corroborated by findings in other plant-insect studies (Charters et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Bell et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Overall, this body of research highlights a complex, multi-layered network where plant genotype, microbial symbiosis, and environmental stress interact to shape herbivore physiology, behavior, and survival. Understanding these dynamics is essential for developing sustainable pest and water management strategies in agroecosystems facing increasing water scarcity.\u003c/p\u003e \u003cp\u003eThe preference of \u003cem\u003eS. avenae\u003c/em\u003e for drought-sensitive wheat cultivars like Xinong-1376 over drought-resistant ones such as Yunhao-618, especially when associated with \u003cem\u003eA. delicata\u003c/em\u003e, aligns with recent findings on plant\u0026ndash;aphid\u0026ndash;AMF interactions under water stress. Similar research highlights that drought-stressed, less resistant cultivars often accumulate higher concentrations of soluble nitrogen compounds and amino acids, improving phloem sap quality and thus aphid nutrition and reproduction (Weldegergis et al. \u003cspan citationid=\"CR123\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Kansman et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The temporal dynamics in aphid colonization, with fluctuations corresponding to plant defense activation and subsequent weakening, reflect complex physiological responses modulated by AMF symbiosis, mirroring observations in other studies where initial plant stress induces defensive compounds that later decline, increasing herbivore susceptibility (Kansman et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Furthermore, research on AMF-mediated drought tolerance shows that while mycorrhizal association can enhance plant resistance, cultivar-specific variability often determines the degree to which aphid colonization is suppressed or facilitated (Volpe et al. \u003cspan citationid=\"CR120\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Sonbol et al. \u003cspan citationid=\"CR108\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). The observed increase in dry and fresh weights of wheat cultivars Xinong-1376 and Yunhao-618 associated with \u003cem\u003eA. delicata\u003c/em\u003e under both well-watered and drought conditions aligns well with recent findings on AMF-mediated plant growth promotion. Similar studies across diverse crops have consistently demonstrated AMF\u0026rsquo;s role in enhancing biomass by improving nutrient and water uptake through an extensive hyphal network, especially under abiotic stress such as drought (Gao et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). For instance, AMF inoculation in tomatoes and cucumbers significantly increased fresh and dry weights, attributed to enhanced photosynthetic efficiency and antioxidant defense that mitigate oxidative stress during water deficit (Alam et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Tang et al. \u003cspan citationid=\"CR113\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). These mechanisms contribute to stabilization of water potential and accumulation of protective metabolites like flavonoids and glycosides (Stallmann et al. 2020). Moreover, cultivar-specific responses to AMF are increasingly recognized, with drought-resistant varieties like Yunhao-618 leveraging AMF symbiosis to sustain growth under stress, consistent with findings in barley and maize (Begum et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Additionally, enhanced root biomass observed with AMF inoculation supports stronger water and nutrient acquisition capacity even in varied drought intensities (Chandrasekaran \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). This reinforces the potential of integrating AMF inoculation into crop management strategies to improve yield stability under fluctuating water availability. Collectively, these studies underscore AMF\u0026rsquo;s broad applicability and cultivar-specific benefits for sustainable agriculture under climate variability.\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eThis study revealed that \u003cem\u003eA. delicata\u003c/em\u003e colonization (62\u0026ndash;67% RLC) significantly enhanced \u003cem\u003eSitobion avenae\u003c/em\u003e performance on both drought-resistant and drought-susceptible wheat cultivars under contrasting water regimes. AMF symbiosis prolonged aphid longevity (13.8 to 24.9 days), increased fecundity (10.2 to 22.4 offspring), and elevated adult body weight (362.6 to 665.40 \u0026micro;g) under water-deficit stress. Enhanced aphid desiccation tolerance was evident through increased fresh body mass, absolute water contents, and reduced water loss rates on AMF-colonized plants. Host preference experiments demonstrated strong aphid attraction to mycorrhizal-associated plants, with population densities reaching 19.20\u0026ndash;23.50 individuals on Xinong-1376 compared to 5.50\u0026ndash;9.30 on Yunhao-618 at 72 hours post-release. AMF colonization significantly improved plant biomass production under both water treatments, confirming beneficial effects on host plant fitness. These results indicate that mycorrhizal-mediated alterations in plant nutritional quality create more suitable conditions for aphid herbivory, potentially offsetting cultivar resistance mechanisms. The tri-trophic interactions observed highlight complex ecological trade-offs where plant-beneficial symbionts may inadvertently enhance pest performance through bottom-up nutritional effects. Cultivar-specific responses suggest genetic variation influences the magnitude of AMF-aphid interactions under environmental stress. These findings emphasize the need to consider belowground symbiotic associations when developing drought-tolerant crop varieties for sustainable pest management. Future research should elucidate the biochemical mechanisms underlying these context-dependent plant-microbe-insect interactions in agricultural ecosystems.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e \u003ch2\u003eCompeting interests:\u003c/h2\u003e \u003cp\u003eThe author declares that there are no competing interests.\u003c/p\u003e \u003c/p\u003e\u003ch2\u003eFunding:\u003c/h2\u003e \u003cp\u003eThe authors did not receive support from any organization for the submitted work.\u003c/p\u003e\u003ch2\u003eAuthor Contributions:\u003c/h2\u003e \u003cp\u003e \u003cb\u003eAGK\u003c/b\u003e: Conceptualization, Data curation, Formal analysis, Methodology, Writing \u0026ndash; original draft. \u003cb\u003eMA\u003c/b\u003e: Writing \u0026ndash; review \u0026amp; editing.\u003c/p\u003e\u003ch2\u003eAcknowledgments:\u003c/h2\u003e \u003cp\u003eWe would like to thank Prof. Yajun Xi from College of Agronomy of Northwest A\u0026amp;F University for providing seeds of different wheat cultivars used in this study.\u003c/p\u003e\u003ch2\u003eData availability:\u003c/h2\u003e \u003cp\u003eThe datasets generated and analyzed during this study are available from the corresponding author upon reasonable request.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAddo-Bediako A, Chown SL, Gaston KJ (2001) Revisiting water loss in insects: a large scale view. 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Annu Rev Plant Boil 68:513\u0026ndash;534. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1146/annurev-arplant-042916-040856\u003c/span\u003e\u003cspan address=\"10.1146/annurev-arplant-042916-040856\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":true,"hideJournal":true,"highlight":"","institution":"Northwest A\u0026F University","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":"Mycorrhiza, aphid, life-history, host choice, water-stress, AMF-plant-aphid interactions","lastPublishedDoi":"10.21203/rs.3.rs-8715853/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8715853/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eAphid-microbe-plant interactions are fundamental to understanding plant responses to biotic and abiotic stressors. Aphid is a significant pest distressing wheat crops globally, especially under drought conditions. The interactions between the aphid \u003cem\u003eSitobion avenae\u003c/em\u003e and arbuscular mycorrhizal fungus (AMF) \u003cem\u003eAcaulospora delicata\u003c/em\u003e, particularly in wheat cultivars under drought conditions, reveal significant dynamics affecting pest performance and plant health. However, the influence of AMF on aphid performance across different wheat-cultivars under water-deficit stress, remains poorly understood. We investigated the effects of AMF on the performance of \u003cem\u003eS. avenae\u003c/em\u003e on two wheat cultivars, Yunhao-618 (drought-resistant) and Xinong-1376 (drought-non-resistant), under varying water conditions. Our results revealed that \u003cem\u003eA. delicata\u003c/em\u003e significantly increased root length colonization in Xinong-1376 (67%) and improves aphid developmental duration, longevity, and fecundity under both well-watered and water-deficit conditions. Notably, aphid nymphs exhibited prolonged DDs without AMF-associations, particularly on Xinong-1376 under drought-stress. In contrast, associations with AMF promoted faster growth rates and higher fecundity in Yunhao-618, suggesting that AMF-association can improve water stress effects. Honeydew production was higher in AMF-associated plants under well-watered conditions. Aphid fresh and dry body masses, and water balance traits tended higher on drought-resistant cultivars Yunhao-618 with AMF association under well-water conditions. Additionally, aphids displayed a preference for AMF-associated drought non-resistant Xinong-1376 plants, indicating a strong influence of AMF on host choice dynamics. Our findings reveal that \u003cem\u003eA. delicata\u003c/em\u003e modulates plant-aphid dynamics by enhancing plant drought resilience while inadvertently promoting aphid fitness. This underscores the complex role of AMF intri-trophic interactions and their broader implications under climate change scenarios.\u003c/p\u003e","manuscriptTitle":"Arbuscular mycorrhizal fungus Acaulospora delicata modulates tri-trophic interactions between wheat cultivars and Sitobion avenae under drought stress","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-01-29 06:19:15","doi":"10.21203/rs.3.rs-8715853/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":"9c7c171d-7f98-42f4-af49-655018762c7b","owner":[],"postedDate":"January 29th, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[{"id":61863291,"name":"Entomology"}],"tags":[],"updatedAt":"2026-01-29T06:19:16+00:00","versionOfRecord":[],"versionCreatedAt":"2026-01-29 06:19:15","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8715853","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8715853","identity":"rs-8715853","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}