Abstract
Aeolian sandy soil occupied one-fifth of the agricultural land in Northeast China; however, effective strategies to mitigate aeolian sandy soil degradation and enhance crop production remain largely unexplored. Here, a two-year field experiment was conducted on the aeolian sandy soil to evaluate the effects of rotational strip peanut/cereal intercropping systems on soil quality and crop yields. The experiment comprised seven treatments: peanut/corn intercropping with strip widths of 3, 6, and 9 m; peanut/millet intercropping with strip widths of 3, 6, and 9 m; and a peanut monoculture serving as the control. We showed that both peanut/corn and peanut/millet intercropping systems significantly enhanced the nutrient levels including total alkaline- hydrolyzable nitrogen, available phosphorus, available potassium, and organic matter in the rhizosphere soil compared with peanut monoculture. Moreover, the activities of soil enzymes such as catalase, cellulase, urease, and phosphomonoesterase were substantially increased in intercropping treatments. Entropy-weighted TOPSIS analysis indicated that peanut/corn and peanut/millet intercropping with strip width of 6-m treatments achieved optimal performance in regulating soil nutrients, enhancing enzyme activities, and improving land-use efficiency. These two treatments also significantly increased the peanut yield and total output values, reduced total input costs, while optimized economic returns over peanut monoculture. In summary, this study provides scientific evidence that rotational peanut/cereal intercropping systems utilizing a 6-m strip width could be utilized as practical strategies for improving crop production, restoring ecological balance, and ensuring sustainable land management of aeolian sandy soil in Northeast China.
Keywords: Intercropping, Aeolian sandy soil, Soil nutrients, Enzyme activities
Introduction
Aeolian sandy soil characterizes the ecologically fragile zones in the arid and semi-arid regions of Northeast China [1]. Aeolian sandy soil adversely affects crop yields due to drought, low fertility, wind erosion, and unstable temperature [2, 3]. For instance, Fuxin City, located at the southern margin of the Horqin sandy land, suffers from low soil fertility and high susceptibility to wind erosion [4]. As the primary oilseed/legume crop in this region, peanut (Arachis hypogaea L.) is predominantly cultivated as a monoculture [5–7]. However, prolonged monoculture can negatively impact soil microbial community structures and deplete nutrient availability, ultimately reducing yields and exacerbating soil degradation [8, 9]. Although corn (Zea mays L.) and millet (Setaria italica L.) are widely cultivated cereal crops in this region which can be integrated into peanut-based cropping systems, research on their rotational strip configurations remains limited.
Intercropping has been extensively considered as crucial agronomic strategies for improving soil properties and maximizing resource utilization efficiency [10, 11]. Literatures have shown that legume/cereal intercropping systems could alleviate the constraints of consecutive monoculture of legume crops and promote the resource utilization efficiency through facilitating interspecific interactions, thereby contributing to the ecological restoration and agricultural productivity enhancement [12–14]. In addition, strip width plays a critical role in regulating interspecific interactions and determining system productivity [15, 16]. In Northeast China, rotational strip intercropping of peanut and maize has been widely adopted as a sustainable agricultural strategy that could increase the contents of alkaline-hydrolyzed nitrogen and available potassium in root zones, and thus sustain the soil resource utilization and boost crop productivity [11, 17–19]. Nonetheless, current research on aeolian sandy-land cultivation patterns has primarily focused on either intercropping or rotation alone, with limited comparisons of rotational strip intercropping systems under different strip widths.
In addition to intercropping, crop rotation has been shown to improve soil nutrient dynamics through multiple mechanisms. The alternating of crop root systems contributes to disrupting compacted soil layers, enhancing the availabilities of soil nutrients, and facilitating soil organic matter accumulation [20–22]. From the yield perspective, monoculture often accelerates soil degradation and yield reduction, whereas rotation serves as an ecologically sustainable approach that maintains productivity while promoting soil health. In a meta-analysis of 11 long-term trials, diversified rotations increased the average corn yields by more than 20% [23]. In another experiment, soybean/corn rotation enhanced the soybean yields by 8–20% and corn yields by 5–30% [24]. Previous studies have also shown that multi-year rotational strip intercropping of peanut and cotton significantly increased cotton yield without reducing peanut yield [25, 26]. Collectively, legume-based rotational strip intercropping system provides a fruitful avenue for farmers to boost crop productivity in aeolian sandy soil.
In the current study, a two-year field experiment was carried out to investigate how rotational strip peanut-based intercropping systems regulate soil nutrients and enzyme activities in aeolian sandy soil. Specifically, we aimed to quantify the effects of varying strip widths (e.g. 3 m, 6 m, and 9 m) within these intercropping systems. The objective of this work was to identify optimal strip widths for rotational peanut/maize and peanut/millet intercropping systems that mitigate continuous monoculture constraints, improve soil quality, enhance economic returns, and ultimately increase crop productivity.
Materials and methods
Plant materials
Corn (Zea mays L., cv. ‘Liaodan 588’) seeds were provided by Corn Research Institute Liaoning Academy of Agricultural Sciences. Peanut (Arachis hypogaea L., cv. ‘Fuhua 30’) seeds were obtained from Liaoning Research Institute of Sandy Land Control and Utilization. Millet (Setaria italica L., cv. ‘Zhongzagu 36’) seeds were purchased from Shanxi Nongwawa Seed Industry Co., Ltd. All of the above crop species are widely cultivated drought-tolerant varieties in this region.
Study area and experimental design
The field study was conducted in 2023 and 2024 at the Zhanggutai Experimental Station (43°20’N, 123°22’E) of the Liaoning Provincial Institute of Sandy Land Management and Utilization, Liaoning, China. The experimental site has a temperate continental monsoon climate, with an annual mean temperature of 7.2℃, mean precipitation of 500 mm, potential evaporation of 1300 mm, and relative humidity of 59%. The dominant soil type is aeolian sandy soil with poor water- and nutrient-retention capacities. The experimental plot has been cultivated with peanut for at least 5 consecutive years prior to this study. The initial soil properties of 0–20 cm soil layer were as follows: pH 6.56, organic matter 8.11 g kg− 1, alkaline nitrogen 23.98 mg kg− 1, available phosphorus 45.40 mg kg− 1, and available potassium 84.86 mg kg− 1. The meteorological data for 2023–2024 is presented in Fig. 1.
Fig. 1.

The average precipitation and temperature for each month from 2023 to 2024 in the experimental site
A total of seven treatments were used: peanut/corn intercropping with strip widths of 3 m (PC1), 6 m (PC2), and 9 m (PC3); peanut/millet intercropping with strip widths of 3 m (PM1), 6 m (PM2), and 9 m (PM3); and consecutive monoculture of peanut (PP). Intercropping was initiated in 2023, with crop rotation adopted in 2024. All crops were planted using ridge cultivation without mulching. Peanut was sown at 50 cm row spacing and 13–15 cm hole spacing, with two seeds per hole. Corn was sown at 50 cm row spacing and 40 cm plant spacing, using single-seed planting. Millet was sown at 40 cm row spacing and 15 cm plant spacing (Fig. 2). All plots received uniform basal application of synthetic fertilizer (750 kg ha-1; N: P2O5: K2O = 1: 1.5: 1.5). No pesticides were used during the entire growing season. Sowing was performed manually in mid-May following standard agronomic procedures: deep furrowing, basal application of fertilizer, soil mounding, sowing, and soil covering with compaction. Irrigation was applied once during the growing period and adjusted based on actual rainfall conditions. The experiment was set up via a complete randomized block design with three biological replicates.
Fig. 2.

Schematic diagram illustrating planting patterns of each treatment from 2023 to 2024
Measurements, data collection, and soil analysis
At late podding stage of peanut, rhizosphere soil samples were collected from peanut, corn, and millet simultaneously in each year. The intact root systems were carefully excavated and the adhered soil was collected using a gentle shaking method to avoid damaging root structures. For each treatment, 15 sampling points were established for each crop species. Each sampling point was evenly distributed across the plot with sufficient spacing to ensure representativeness. Then, soil from 5 sampling points was combined to form a composite sample. For intercropping treatments, the composite samples from two species were randomly combined to form one replicate. The samples were liquid frozen immediately and delivered to the laboratory. After which, visible impurities such as plant residues and stones were removed, and the samples were air-dried in a ventilated environment. After drying, the soil samples were passed through a 1 mm sieve and divided into two subsamples: one stored at 4℃ for enzymatic assays and the other stored in a cool, dry location for nutrient analysis.
Soil nutrient analyses were performed following the Soil Agrochemical Analysis Protocol with minor modifications [27]. Soil total alkali-hydrolyzable nitrogen (AHN) was determined with the alkaline hydrolysis diffusion method. Available phosphorus (AP) was assessed by the NaHCO3 extraction method. Available potassium (AK) was analyzed by neutral ammonium acetate extraction followed by flame photometry. Finally, soil organic matter (SOM) analysis was carried out using the potassium dichromate, dilution-heat oxidation method.
Soil catalase activity was determined volumetrically by mixing the soil with distilled water and H2O2, shaking for 20 min, adding 1.5 mol L− 1 H2SO4, filtering 25 mL of solution, and titrating with potassium permanganate until a pale pink endpoint. Cellulase activity was evaluated using the 3, 5-dinitrosalicylic acid colorimetric method. Urease activity was also analyzed using the sodium hypochlorite colorimetric method. Phosphomonoesterase activity was assessed using the p-Nitrophenyl phosphate method [28].
Crop yield and economic benefits
Crop yield
During crop maturity, three plots of each crop were randomly selected within each treatment, including one consecutive monoculture of peanut plot and relevant plots in intercropping and rotation treatments. Harvested yields were converted to per-hectare values based on actual harvested area.
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(2)
Land use efficiency
Land use efficiency was evaluated using the crop land equivalent ratio (LER), calculated with consecutive monoculture yield as the control:
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1 |
where Yc denotes yield under consecutive monoculture and Yr denotes yield under rotation.
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(3)
Economic benefit analysis
Economic benefit parameters were calculated using production input data (seeds, fertilizers, pesticides, labor, and machinery) from field trials (2023–2024) for corn, peanut, and millet. Crop procurement prices were obtained from official platforms such as the Liaoning Provincial Department of Agriculture and Rural Affairs and the Grain and Material Bureau, complemented by local market prices in Zhangwu County. Economic indicators were calculated as follows:
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2 |
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3 |
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4 |
Statistical analysis
Data were expressed as means ± standard deviations. Statistical analyses was performed using Microsoft Excel and SPSS 26.0. Data were tested for normality and homogeneity of variance before parametric analysis. One-way and two-way ANOVA followed by least significant difference (LSD) test were utilized to assess significant treatment differences at P < 0.05, P < 0.01, and P < 0.001. Data visualization was conducted using Origin software. A Partial Least Squares Path Model (PLS-PM), implemented using the plspm package in R, was applied to evaluate the relationships among strip width, soil nutrient contents, soil enzyme activities, crop yields, and land use equivalent ratio.
Results
Our first objective was to evaluate the impact of peanut-based rotational strip intercropping systems on soil nutrient accumulations. Relative to intercropping in 2023, all intercropping treatments in 2024 exhibited significant increases in AHN, AP, and AK by 10.40-31.75%, 35.99–95.15%, and 16.61–60.29%, respectively. In contrast, consecutive monoculture of peanut (PP) resulted in a 42.93% decline in AHN, although AP and AK rose by 47.35% and 25.67%, respectively. For SOM, all intercropping treatments except PC3 showed increases ranging from 7.80% to 47.43%, while PP also exhibited a moderate increase of 14.71% (Fig. 3). Overall, intercropping systems, particularly PC3, PC2, and PM2, consistently outperformed consecutive monoculture of peanut in most nutrient metrics including AHN, AP, AK, and SOM, demonstrating pronounced improvements in nutrient accumulations in the rhizosphere soil.
Fig. 3.

Nutrient contents of rhizosphere soil under different treatments
The effect of peanut-based rotational strip intercropping systems on soil enzyme activities was further elucidated. Across intercropping systems, the activities of all four measured soil enzymes-catalase, cellulase, urease, and phosphomonoesterase-showed an overall increase from 2023 to 2024. Compared with 2023, soil catalase activity significantly increased by 12.47–71.41% under rotation systems, whereas PP showed only a marginal increase of 12.47%. In 2024, soil cellulase activity exhibited the most pronounced response under rotation treatments, with PP still showing a considerable rise. Similarly, soil urease activity was significantly increased in 2024 in all treatments, compared with 2023. Moreover, soil phosphomonoesterase activity rose by 16.67–47.52% across rotation treatments and PP still displayed a 27.47% increase. Overall, in 2024, peanut/corn and peanut/millet rotation systems markedly enhanced the activities of catalase, cellulase, and urease, compared with PP with a few exceptions. However, no significant changes were observed between intercropping treatments and PP in soil phosphomonoesterase activity of 2024 (Fig. 4).
Fig. 4.

Soil enzyme activities in the rhizosphere under different treatments
Averaged for two years, the peanut yield under different treatments followed the order PM2 > PC2 > PM3 > PC3 > PC1 > PM1 > PP. Compared with PP, these treatments significantly increased the peanut yield by 16.43%, 42.71%, 30.45%, 9.31%, 44.16%, and 32.09%, respectively. Based on planting strip width, the yield-promoting effect was most pronounced at 6 m, followed by 9 m and 3 m. Similar trends were observed for corn and millet, where yields also ranked as 6 m > 9 m > 3 m, indicating that a 6-m strip width seems to be the most effective intercropping/rotation configuration in this experiment. The crop land equivalent ratio (LER) further showed that all peanut LER values exceeded 1, except for PM1, demonstrating that rotational strip peanut-based intercropping systems displayed obvious intercropping advantages over peanut consecutive monoculture system (Fig. 5).
Fig. 5.

Coefficients of annual average yield and land equivalent ratio of each crop under different treatments
The economic benefit analysis for 2023–2024 incorporating input cost, total output value, and net return value was further carried out. Although input costs varied only slightly among treatments (approximately 1,530-2,980 $ ha− 1), substantial differences were observed in output values and net returns due to crop-type and yield variations. In particular, PP generated the lowest net income (1,960 $ ha− 1), whereas the PC2 treatment achieved the highest output value (9,993 $ ha− 1) and net return (7,012 $ ha− 1), representing a 2.5-fold increase compared with PP and indicating superior economic performance. The PC1 and PC3 treatments followed, yielding net returns ranging from 4,670 to 6,205 $ ha− 1. For PM2, the average annual input cost was 2,934 $ ha− 1, with a total output value of 8,913 $ ha− 1 and a net return of 5,979 $ ha− 1, 67.23% higher than PP (Fig. 6). These results confirm that rotational strip peanut-based intercropping systems substantially enhance economic returns compared with PP. Moreover, intercropping strip width plays a decisive role, with a 6-m strip width producing the most favorable economic returns.
Fig. 6.

Economic benefit analysis of each treatment from 2023 to 2024
PLS-PM analysis further clarified the relationships among strip width, soil nutrients, soil enzyme activities, crop yield, and LER. In peanut/corn intercropping system, strip width exerted a highly significant positive influence on LER and significantly enhanced soil nutrient levels. Soil enzyme activity also showed significant positive effects on both soil nutrients and yield, yet displayed a highly significant negative effect on LER. Moreover, yield contributed positively and significantly to LER. In the peanut/millet intercropping system, strip width significantly increased both yield and LER, but exerted a significant negative effect on soil nutrients. Soil enzyme activity strongly and positively influenced both soil nutrients and yield. Again, yield showed a highly significant positive impact on LER (Fig. 7a & b). Standardized total effects indicated that both strip width and yield were strong drivers of LER in peanut-based intercropping systems, whereas strip width showed the greatest overall influence on the peanut/millet intercropping system (Fig. 7c & d).
Fig. 7.

Partial Least Squares Path Models (PLS-PM) for rotational peanut/corn intercropping (a) and rotational peanut/millet intercropping (b) systems. Standardized total effects of strip width, soil enzyme activities, soil nutrients, and yield on land equivalent ratio in rotational peanut/corn intercropping (c) and rotational peanut/millet intercropping (d) systems. Observed or latent variables are illustrated in boxes. One thousand bootstrap resamples were conducted to estimate the path coefficients. Positive and negative effects are represented by black and red arrows, respectively. Path coefficients that are not significantly different from zero are shown as grey dashed lines; *P < 0.05, **P < 0.01, and ***P < 0.001. Percentages above the boxes indicate the explanatory power (R²) of the variables. Model fit was assessed using the goodness-of-fit (GoF) index
Discussion
Intercropping combined with crop rotation represents an efficient agronomic strategy for mitigating soil degradation and enhancing crop yields [29–31]. The underlying mechanisms driving these benefits include: (1) improve light utilization via canopy stratification, (2) optimize nutrient distribution through modifying root architecture and enhancing nutrient cycle, and (3) mitigate pests and diseases by leveraging crop resistance variation [32–34]. The current 2-year field study carried out in aeolian sandy soil provides evidence that rotational strip (strip width of 6 m) peanut/corn and peanut/millet intercropping systems exhibited significant higher levels of nutrients including AHN, AP, AK, and SOM, as well as crucial enzymes in the rhizosphere soil, and ultimately increased the crop yields compared with consecutive monoculture of peanut. The findings of this study demonstrate that rotational strip peanut/cereal intercropping can serve as a novel strategy to improve soil properties and crop productivity in aeolian sandy soils.
Rotational strip intercropping treatments increased AHN from 2023 to 2024, whereas AHN in PP was dramatically reduced (Fig. 3a), which was consistent with the earlier reports that rotational strip legume/cereal intercropping systems promote soil nitrogen accumulation and availability [35, 36]. These effects may result from improved soil environment and microbial community structure [37] as well as enhanced legume nodulation and nitrogen fixation facilitated by their neighboring crops, which increases nitrogen retention [38]. In contrast, fungal overgrowth and microbial depletion might be the crucial reason for the reduction of nitrogen availability under consecutive monoculture of peanut [39, 40]. Additionally, the higher AP content under rotational strip intercropping treatments aligns with the improvement of soil micro-ecological environment, compared with consecutive monoculture of peanut (Fig. 3b). The above findings were consistent with previous reports which demonstrate that enhanced microbial activities in rhizosphere soil under legume-based intercropping treatments could facilitate soil organic phosphorus mineralization, optimize phosphorus allocation and partition to productive organs, and consequently sustain higher phosphorus use efficiency [41–43]. In contrast, consecutive monoculture of peanut suppresses microbial function and therefore reducing AP availability. Similarly, AK content increased under rotational strip intercropping treatments compared with consecutive monoculture of peanut (Fig. 3c). Consistent with the previous studies, we demonstrate that legume-cereal intercropping could help maintain the stability of the soil microenvironment and facilitation root nutrient absorption [44–46]. Moreover, intercropping systems increase the crop diversity, thus promote potassium residue decomposition release; however, peanut monoculture restrict these processes. Again, the soil organic matter accumulation was also greater under rotational strip intercropping systems (Fig. 3d). This phenomenon can be explained by the mechanism that legume crops could improve soil structure, accelerate decomposition and transformation of soil organic matter, and maintain carbon turnover balance in legume-based intercropping systems [47–49]. Conversely, consecutive monoculture promotes soil compaction that impedes soil organic matter buildup.
Soil enzyme activities are core biological indicators for assessing soil health and ecological functions due to their high sensitivities to environmental changes [50, 51]. Soil enzymes also serves as key biological drivers in the formation of soil fertility for they directly drive the nutrient transformation processes in the rhizosphere soil [52, 53]. The induction of catalase activities in PC and PM treatments (Fig. 4a) indicate that rotational strip intercropping facilitates soil biological oxidation processes and contributes to the maintenance of material and energy transformation efficiencies [54]. Cellulase catalyzes the degradation of cellulose into monosaccharides such as glucose, providing energy substrates and carbon sources for organisms, thereby maintaining the balance of the soil carbon cycle [55]. We therefore speculate that the induction of cellulase activities in PC and PM treatments (Fig. 4b) promote the mineralization of plant residues which release available nutrients into the soil, thus regulates soil fertility and supports plant growth. Urease, as a core catalytic enzyme in the nitrogen cycle, directly indicates the decomposition efficiency of nitrogen-containing organic matter in soil [56]. Thus, the elevated urease activity in the PC and PM treatments (Fig. 4c) contributes to the precise regulation and effective supply of nitrogen in the rhizosphere. Soil phosphatase helps to provide phosphorus to microorganisms which activity directly indicates the efficiency of soil phosphorus transformation [57]. Therefore, it is tempting to speculate that the induction of phosphatase activities in PC and PM treatments (Fig. 4d) promote the mineralization of organic phosphorus to release available phosphorus in peanut-based intercropping systems.
Notably, peanut-based intercropping systems with moderate strip width of 6 m significantly increased crop yields compared with other treatments (Fig. 5). On the one hand, moderate strip width could prevent the intensified light competition between the intercropped species (primarily affecting the mid-to-lower canopy crop, such as peanut) that occurs in narrower configurations like the 3-m strip width. On the other hand, a moderate strip width prevents the attenuation of interspecific interactions caused by excessively wide strips such as 9 m. Therefore, 6 m was taken as the optimum strip width in peanut-based intercropping systems that could: (1) maintain the canopy microenvironment parameters such as light intensity, temperature, humidity, and CO2 concentration in a relatively optimal range; (2) promote the belowground interspecific facilitation by utilizing interspecies niche differentiation and root spatial complementarity. Additionally, PLS-PM analysis also indicate a strong relationship between strip width and LER where LER calculations confirmed synergistic yield benefits across rotational strip intercropping systems (Fig. 7).
Conclusions
In conclusion, we demonstrate that rotational strip peanut/corn and peanut/millet intercropping systems with a 6-m strip width simultaneously enhanced crop yields and economic returns by optimizing interspecific interactions, as evidenced by the promotion of nutrient levels (total alkaline- hydrolyzable nitrogen, available phosphorus, available potassium, and organic matter) and enhanced activities of enzymes (catalase, cellulase, urease, and phosphomonoesterase) in the rhizosphere soil. These results provide a practical framework for the sustainable utilization of aeolian sandy soil in Northeast China and offer quantitative guidance for agricultural system reform in similar ecologically fragile regions. To address the limitation of the two-year duration, extended multi-year field experiment is still required to further elucidate the long-term rotational effects of peanut/cereal intercropping systems.
Abbreviations
- AK
Available potassium
- ANOVA
Analysis of variance
- AP
Available phosphorus
- LER
Land equivalent ratio
- LSD
Least significant difference
- PLS-PM
Partial least squares path model
- SOM
Soil organic matter
- AHN
Alkali-hydrolyzable nitrogen
Authors’ contributions
PS, XZ, and TS conceived and designed the experiments. JL, HW, QT, YW, GY, and WB performed the experiments. TS contributed to the writing of the manuscript. All authors read and performed the final editing of the manuscript.
Funding
This study was financially supported by the Applied Basic Research Program of Department of Science & Technology of Liaoning Province (2022020385-JH2/1013) and the Demonstration and Extension of Integrated Technologies for High-Yield, High-Quality, and High-Efficiency Peanut Production of Department of Agriculture & Rural Affairs of Liaoning Province (2026-NZZY-05).
Data availability
All data generated or analyzed during this study are included in this published article.
Declarations
Ethics approval and consent to participate
All field work associated with this research was performed in accordance with the laws and guidelines of Liaoning Province and the People’s Republic of China.
Consent for publication
All authors have read and approved the final manuscript for publication.
Competing interests
The authors declare no competing interests.
Footnotes
Publisher’s note
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Puxiang Shi and Jing Li contributed equally to this work.
Contributor Information
Xiaoxia Zou, Email: xxzou@qau.edu.cn.
Tong Si, Email: tongsi@qau.edu.cn.
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Associated Data
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Data Availability Statement
All data generated or analyzed during this study are included in this published article.




