Adaptation of Asparagus munitus to karst adversity and its effects on soil

Adaptation of Asparagus munitus to karst adversity and its effects on soil | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Article Adaptation of Asparagus munitus to karst adversity and its effects on soil Mingsheng Zhang, liu tang, miao liu, liyang shangguan, haidong Huang, and 2 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-4318709/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 In this study, Asparagus munitus was planted in karst rocky desertification (KRD) areas and arable land (AL) to explore the changes in plant adaptability to KRD environments and effects on KRD soils. Results showed A. munitus exhibited shrinkage in KRD habitats, evidenced by increased number and length of stems, branch length, and other agronomic traits; significantly elevated antioxidant enzyme and osmoregulator content in leafy shoots; and increased polysaccharide content in tuberous roots. Correlation and principal component analyses of plant agronomic traits revealed strong correlations among indicators except basal stems, with leaf shoot length and root diameter as the first and second principal components, respectively. Soil physical properties and nutrient contents showed KRD soil pH, water content, porosity, and nutrient content markedly changed after A. munitus planting, overall positively developing. This study reveals partial adaptive strategies of A. munitus to KRD habitats and ameliorative effects of planting A. munitus on KRD soils, providing theoretical bases for A. munitus cultivation and KRD soil improvement. Biological sciences/Physiology Biological sciences/Plant sciences Earth and environmental sciences/Environmental sciences Asparagus munitus karst rocky desertification agronomic traits soil physical properties soil nutrients Introduction According to the latest karst rocky desertification (KRD) survey, China's KRD area was 7.22 million hm2 in 2021 [ 1 ] . KRD severely limits human land use due to poor nutrient storage, severe temporary drought, and poor ecological stability [ 2 ] . Studies have shown a correlation between poverty and the KRD environment [ 3 , 4 ] . To address ecological and economic issues, controlling KRD is urgently needed for environmental protection and poverty eradication. Asparagus cochinchinensis (Lour.) Merr. is a perennial herb; its tuberous roots have long been used in traditional Chinese medicine and was included in the 2020 Chinese Pharmacopoeia [ 5 ] . Currently, Asparagus munitus Wang et S.C. Chen ( A. munitus ), Asparagus lycopodineus (Baker.) Wang et. Tang ( A. lycopodineus ) and Asparagus subscandens Wang et S.C. Chen ( A. subscandens ) have been artificially domesticated [ 6 ] . Although only A. cochinchinensis is officially recognized, the other three Asparagus species are more widely cultivated due to A. cochinchinensis 's low yield and their similar medicinal composition. With Asparagus development and utilization, there are great market prospects in pharmacology [ 7 – 10 ] and health foods [ 11 ] . A. munitus 's wide distribution indicates strong environmental adaptability [ 12 ] . Several studies have shown that herbaceous plants undergo better carbon sequestration than woody plants [ 13 ] , and their well-developed root systems are able to fix the topsoil layer and prevent soil erosion [ 14 ] . The effects of planting Asparagus on soil physicochemical properties and nutrients have been studied [ 15 ] , but few studies exist on Asparagus planting in karst soil and its interaction. This study compared A. munitus changes in KRD and arable land (AL) to predict survival strategies in KRD. It also tested whether A. munitus could improve KRD soil conditions, providing a basis for its cultivation and KRD control and new ideas for medicinal plant development. Results A. munitus under KRD conditions shows a decreasing trend The results revealed significant differences (p < 0.05) in stem number, stem length, middle lateral branch length and leaf length between the EG and CK plants of A. munitus (Table 1 ). No significant differences were observed for the other indicators. LSD indicated no significant difference in below-ground biomass, however CK averaged approximately one-third higher than EG, EG tubers tended to be shorter but greater in diameter (Fig. 2 A, B). Overall, EG growth was inhibited, especially above ground, and was weaker than that in the CK treatment. Table 1 Agronomic traits of A. munitus plants grown in different regionsa and b indicate significant differences at the 0.05 level ( p < 0.05); A and B indicate significant differences at the 0.01 level (p < 0.01). The values are the means ± standard deviations Trait Unit Control check Experiment group Number of stems each 5.33 ± 0.82 A 3.67 ± 0.52 B Basal stem thickness mm 3.05 ± 0.46 a 2.98 ± 0.53 a Stems length cm 141.66 ± 16.8 a 114.05 ± 21.0 b Lateral branch length cm 21.97 ± 2.84 A 16.85 ± 1.88 B Leafy shoot length mm 6.76 ± 0.87 A 5.18 ± 0.61 B Above-ground biomass g 42.30 ± 10.19 a 34.88 ± 6.64 a Underground biomass g 245.53 ± 62.67 a 186.76 ± 30.11 a Biochemical responses of A. munitus leafy shoots to KRD The stomatal density in the EG treatment was significantly greater than that in the CK treatment (Fig. 3 ), averaging 471.26 mm − 2 . Compared to those in the CK treatment, the photosynthetic pigment content in the EG treatment was markedly lower; however, the carotenoid-to-chlorophyll ratio differed highly significantly between the groups (Table 2 ). Similarly, the leaf SOD, POD, and CAT activities in the EG treatment were 1.4, 1.6 and 1.7 times greater than those in the CK treatment, respectively. Osmoregulatory substances showed similar increasing trends in the EG compared to the CK. These results suggest that A. munitus individuals experience some degree of stress under KRD conditions. Table 2 Biochemical indicators of plants in different environments. A and B indicate significant differences at the 0.01 level ( p < 0.01). The values are the means ± standard deviations. FW: fresh weight Trait Unit Control check Experiment group Stomata density Pcs/mm 2 375.43 ± 45.92 B 471.26 ± 103.52 A Chlorophyll a mg/g FW 1.81 ± 0.03 A 1.43 ± 0.04 B Chlorophyll b mg/g FW 0.72 ± 0.058 A 0.52 ± 0.033 B Carotenoid mg/g FW 0.41 ± 0.002 A 0.37 ± 0.004 B Carotenoids/chlorophyll ×10 − 2 16.27 ± 0.12 B 18.83 ± 0.49 A The medicinal components of A. munitus are not affected by KRD Total saponins, polysaccharides and total amino acids are the main medicinal components of A. munitus tubers. The results showed that the polysaccharide content was significantly greater ( p < 0.05) in the EG tubers than in the CK tubers, while no significant differences were observed for the other components (Fig. 4 ). These findings indicate that A. munitus plants can accumulate medicinal components normally when planted in KRD areas, which is even slightly higher than what is observed in the CK treatment. Correlation analysis and principal component analysis of agronomic traits in A. munitus Correlation analysis revealed strong associations between most agronomic traits, except for basal stem thickness, which was less correlated with the other indicators (Fig. 5 ). After removing the basal stem thickness, the remaining traits were subjected to principal component analysis. The first two principal components had eigenvalues above 1 (5.248 and 1.424) and accounted for 83.400% of the total variance (Table 3 ). Thus, these two components represent the majority of the information from the eight agronomic traits. Leafy shoot length and tuber diameter had the highest eigenvector values for the first and second principal components (Table 4 ). Therefore, the principal components included leafy shoot length and tuber diameter. Table 3 Principal component analysis of the agronomic traits of A. munitus Components Eigenvalue Contribution rate Cumulative contribution rate 1 5.248 65.594 65.594 2 1.424 17.806 83.400 3 0.513 6.411 89.811 4 0.343 4.289 94.100 5 0.258 3.221 97.320 6 0.119 1.485 98.806 7 0.077 0.968 99.773 8 0.018 0.227 100.000 Table 4 Principal component analysis eigenvectors and contributions of agronomic traits Agronomic trait Eigenvector Principal Component 1 Principal Component 2 Number of stems 0.867 -0.198 Stems length 0.782 0.321 Lateral branch length 0.851 0.160 Leafy shoot length 0.976 -0.010 Above-ground biomass 0.793 0.540 Underground biomass 0.951 0.083 Tuber length 0.821 -0.228 Tuber diameter -0.025 0.951 Table 5 Effect of A. munitus planting on the physical properties of KRD soils. a and b indicate significant differences at the 0.05 level ( p < 0.05); A and B indicate significant differences at the 0.01 level ( p < 0.01). The values are the means ± standard Trait Unit Soil control Soil near A. munitus pH 7.37 ± 0.05 A 7.21 ± 0.06 B soil water content % 24.50 ± 1.10 B 30.76 ± 0.97 A soil bulk density % 0.93 ± 0.02 b 1.02 ± 0.04 a soil total porosity % 59.81 ± 4.24 B 62.15 ± 0.92 A soil capillary porosity % 54.83 ± 1.48 b 58.78 ± 0.89 a soil noncapillary porosity % 2.64 ± 0.21 B 3.37 ± 0.12 A Table 6 Effect of A. munitus planting on the soil nutrient content of KRD soils. a and b indicate significant differences at the 0.05 level ( p < 0.05), and A and B indicate significant differences at the 0.01 level ( p < 0.01). The values are the means ± standard deviations Trait Unit Soil control Soil near A. munitus Soil organic matter % 5.80 ± 0.23 B 8.09 ± 0.40 A Total nitrogen g/kg 3.15 ± 0.17 B 4.04 ± 0.21 A Total phosphorus g/kg 0.85 ± 0.04 a 0.72 ± 0.05 b Total potassium g/kg 12.82 ± 0.12 a 11.07 ± 0.54 b Alkali-hydro nitrogen g/kg 243.87 ± 10.04 B 317.72 ± 13.14 A Available phosphorus mg/kg 6.26 ± 0.75 A 2.32 ± 0.19 B Available potassium mg/kg 119.33 ± 10.02 a 94.33 ± 6.43 b A. munitus Planting Changes the Soil Physical Properties in the KRD Area The KRD test site was weakly alkaline. Planting A. munitus significantly reduced the SA pH by 0.16 units on average compared to that in SC ( p < 0.01). Additionally, SA had an extremely significantly higher SWC versus SC ( p < 0.01), at more than 31%. SBD, SP, SCP and SNP all differed significantly between SA and SC to varying degrees. These results demonstrated that A. munitus strongly affects the basic physical properties of KRD soils. Planting A. munitus altered the KRD soil nutrient content The KRD soil nutrient levels varied according to fertility grade. SOM and TN were enriched (Class I), AK was moderate (Class III), and AP was depleted (Class IV), indicating that the site was relatively deficient in P and K. Planting A. munitus significantly increased SOM, TN and AN in SA versus SC ( p < 0.05). TP and AP differed significantly between SA and SC, with SC having 2.7 times greater AP. TK and AK were also significantly greater in the SC treatment than in the SA treatment ( p < 0.05). Taken together, these findings demonstrated that A. munitus planting depletes soil P and K, especially P. Overall, A. munitus decreased the KRD soil pH but increased the SWC, SBD, SP, SOM, TN and AN while decreasing P and K. These data indicate that planting A. munitus significantly alters basic physicochemical properties of KRD soil, largely in beneficial ways. Discussion Adaptation of A. munitus to KRD habitats To adapt to the harsh conditions of KRD habitats, plants exhibit a range of altered morphological and physiological characteristics [ 16 ] . A. munitus exhibited shortened stems, branches, and leaves and a reduced aboveground biomass in KRD areas, indicating an overall "shrinkage" phenotype. This finding aligns with that of other KRD-adapted plant species [ 17 ] and may be linked to the seasonal drought commonly experienced in these areas [ 18 ] . The significantly greater stomatal density observed in the EG mirrors similar adaptive increases under drought conditions reported in Berberis poiretii [ 19 ] . By reducing the number of leafy shoots and the aboveground light-exposed area while increasing stomatal transpiration per unit area, A. munitus may be minimizing water loss and heat damage. This shrinkage strategy could aid survival in the water-limited KRD habitats. Compared to those in the CK treatment, plants in the EG treatment had significantly lower total chlorophyll but a greater carotenoid-to-chlorophyll ratio. This finding aligns with that of other karst species under drought stress [ 20 ] and may be attributed to the photoprotective role of carotenoids in addition to their functions in light absorption and energy transfer [ 21 ] . Such adaptations enable the plants to thrive in the challenging KRD environment. The results of the antioxidant enzyme activity and osmoregulatory substance content analyses showed that plants were subjected to a certain degree of adverse stress in the KRD habitat [ 22 ] . Tests of the medicinal properties of the harvested tubers showed that the polysaccharide content in the EG treatment was significantly higher than that in the CK treatment, which could be related to the physiological functions of polysaccharides [ 22 – 24 ] . Correlation analysis and principal component analysis of the agronomic traits of A. munitus revealed strong correlations among most agronomic indicators except for basal stems. These results could be simplified into two principal components, leafy shoot length and tuber diameter, which was similar to the results of a previous study [ 25 ] , suggesting that the belowground portion of A. munitus may play a more vital role in the plant's growth and development. Planting A. munitus to improve KRD soil conditions The results of the study revealed that the planting of A. munitus significantly reduced the soil pH and augmented the SWC, providing a favorable environment for plant survival. However, the SBD was greater in the SA treatment group than in the control group, unlike previous findings [ 26 ] . This may be due to the high gravel and stone contents at the sampling site affecting the results. SP, SCP and SNP were significantly greater in SA than in SC, which was potentially related to root growth or increased SOM improving soil structure and looseness [ 27 ] . SOM, TN and AN were significantly higher in SA than in SC. SOM is central to soil fertility and vegetation recovery [ 28 , 29 ] . Related studies have shown that N limits early-mid-stage karst ecosystem recovery [ 30 ] . Fern rhizosphere soils have lower AP than non-rhizosphere soils [ 31 ] , but the difference here was greater. This could be because of the already low substrate P concentration coupled with the high P demand during A. munitus growth. Similarly, low soil P in A. munitus habitats was previously observed [ 12 ] , suggesting that P depletion is linked to high P requirements and needs further verification. Overall, A. munitus decreased the KRD soil pH but increased the SWC, SBD, SP, SOM, TN and AN while decreasing P and K. The data indicate that planting A. munitus significantly alters the basic physical and nutritional properties of KRD soil, largely in beneficial ways. The results of the study revealed that the planting of A. munitus significantly reduced the soil pH and augmented the SWC, providing a favorable environment for plant survival. However, an unexpected outcome was noted in the SBD, which was more pronounced in the treatment group with A. munitus (SA) than in the control group, deviating from the findings of Peng et al [ 26 ] . This anomaly could be attributed to the high concentration of gravel and stones at the sampling site, which might have influenced the results. The SP, SCP, and SNP were markedly higher in SA than in the control group, which was potentially associated with enhanced root growth or improved soil structure and looseness as per Viviana et al [ 27 ] . The SOM, TN, and AN were significantly elevated in SA compared to SC. SOM is pivotal for soil fertility [ 28 ] and vegetation recovery [ 29 ] . Related studies have revealed that nitrogen constraints early-mid-stage karst ecosystem recovery [ 30 ] . Fern rhizosphere soils exhibit lower AP than non-rhizosphere soils [ 31 ] , but the difference observed in this study was more pronounced. This could be attributed to the already low substrate P concentration coupled with the high P demand during A. munitus growth. Similarly, low soil P in A. munitus habitats was previously observed [ 12 ] , suggesting that P depletion is linked to high P requirements and necessitates further verification. In summary, A. munitus reduced the KRD soil pH but increased the SWC, SBD, SP, SOM, TN, and AN while decreasing P and K. The data underscore that planting A. munitus significantly alters the basic physical and nutritional properties of KRD soil, predominantly in beneficial ways. Conclusion In conclusion, this experiment initially confirmed that A. munitu could thrive in the KRD environment. This was evident from the morphological changes and physio-chemical reactions that occurred in the plant and its surrounding environment. After planting A. munitu , the physical and chemical properties of KRD soil changed, with increased soil water holding capacity and overall fertility, making it more suitable for other plants. Planting A. munitu in KRD areas not only brings economic benefits but also improves KRD soil conditions and ecological value. This lays a good foundation for KRD management and sustainable development in karst areas. Materials and methods Test site and plant materials The test area is located at the experimental base (26°25′54″N, 106°40′53″E) of Guizhou University, Huaxi District, Guiyang City, Guizhou Province, China. This area features a karst plateau landform with a humid subtropical monsoon climate, an average elevation of 1178 m, an average annual temperature of 20.21°C, and an average annual precipitation of 1,318.90 mm. The vegetation is mainly natural secondary forest. The germplasm resources of A. munitus were sourced from the Planting Base of Asparagus in Xingren Town of Qiandongnan (in Guizhou Province, China), and were identified by Prof. Zhang Mingsheng of Guizhou University. The plants were sampled on April 20, 2023 (Fig. 1 ). The effect of karst environments on A. Munitus. Measurement of agronomic traits A. munitus plants were randomly selected from the KRD habitat as the experimental group (EG), and plants from the AL habitat as the control (CK). The number of plant stems, basal stem thickness (mm), stem length (cm), lateral branch length (cm), foliage branch length (mm), tuber length (cm) and diameter (cm) were measured. an electronic balance was used for biomass measurements (the dry parts and sediment attached to the plants were removed, and the plants were subsequently dried in an electric oven at 60°C until the weight was constant). Measurement of physio-biochemical indicators The leafy shoots of A. munitus were randomly collected and rapidly frozen in liquid nitrogen to the laboratory. The stomata density was calculated according to the methods of Ming and Chen [ 32 ] . The chlorophyll content in the leafy shoots was measured according to the methods of Li [ 33 ] . Superoxide dismutase (SOD), peroxidase (POD), catalase (CAT), soluble sugar, and proline were determined by using a kit (Suzhou Keming Biotechnology Co., Ltd., China), and the soluble protein content was determined by the Caulmers Brilliant Blue method with reference to Li [ 33 ] . Measurement of medicinal components A. munitus tubers were collected randomly from two habitats, boiled in boiling water for 10 min, peeled, and subsequently dried at 60°C and crushed. The total saponin, total amino acid and polysaccharide contents of the tubers were determined by the methods of Li et al. [ 34 ] and Wu Fei et al [ 35 ] . Measurement of soil physical properties On several consecutive sunny afternoons, six soil samples near A. munitus plants (SA) in the KRD area and unplanted blank controls (SC) were collected to test physicochemical properties using methods in "Soil Agrochemical Analysis" [ 36 ] . Soil pH was determined by electrode method. SA and SC soils from 0–20 cm were collected by ring knife method, and assessed for soil bulk density (SBD), soil water content (SWC), soil total porosity (SP), soil capillary porosity (SCP), and soil non-capillary porosity (SNP) using the following formulae: M (ring knife mass), M 1 (soil fresh mass), M 2 (soil saturated mass), M 3 (soil resting mass for 12 h), M 4 (soil air-dried mass), and V (ring knife volume). Measurement of soil nutrients Soil organic matter (SOM) content was determined via potassium dichromate oxidation, total nitrogen (TN) via Kjeldahl method, total phosphorus (TP) via HClO 4 -H2SO 4 , total potassium (TK) via sodium hydroxide fusion and flame photometry, alkali-hydro nitrogen (AN) via alkaline diffusion, available phosphorus (AP) via sodium bicarbonate extraction, and molybdenum antimony anti-colorimetry, and available potassium (AK) via flame photometry method [ 36 ] . The level of soil nutrient grade was evaluated according to the method of Hu et al [ 37 ] . Declarations Author Contribution MS Z did supervision. LT and ML designed and conducted the experiment. LT, L S, and Y H performed collection of data and laboratory analysis. LT and M L performed statistical analysis of the data. LT, GG L and HD H were involved in writing––review and editing of manuscript. All authors have read and approved the final version of manuscript. Data Availability The datasets used and/or analyzed during the current study are available from the Liu Tang on reasonable request. Collection license statement for Cremastra appendiculata Statement: The plant materials used in this study have been obtained with the necessary permits or licenses, complying with all applicable laws and regulations. The plant materials have been formally identified by Professor Mingsheng Zhang at Guizhou University and are deposited in the specimen repository at Guizhou University for reference and verification. Statement of Compliance with Endangered Species Research Policies We hereby affirm our commitment to adhere to relevant institutional, national, and international guidelines and legislation when conducting experimental and field research on wild plants. Specifically, we pledge to comply with the IUCN Policy Statement on Research Involving Species at Risk of Extinction and the Convention on the Trade in Endangered Species of Wild Fauna and Flora. We will respect principles of plant material collection and ensure strict adherence to ethical standards and best practices in the collection, use, and handling of plant samples. We will endeavor to minimize our impact on plant ecosystems in our research and mitigate any potential negative effects on endangered species. We will collaborate with local and international conservation organizations to ensure that our research does not result in overexploitation, misuse, or destruction of plant species. We will promote the conservation and sustainable use of plant biodiversity to support ecosystem health and human well-being. We solemnly pledge that our research will uphold the highest scientific standards and ethical principles to contribute positively to the protection of natural environments and biodiversity. Declaration of interests ☑ The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper. ☐ The authors declare the following financial interests/personal relationships which may be considered as potential competing interests: References Ning, X. B. et al. Main Results of the Fourth Rocky Desertification Survey in Karst Area and Analysis of Rocky Desertification Control Situation. For. Resour. Manage 3, 09–14, http://dx.doi.org/10.13466/j.cnki.lyzygl.2023.03.002 (2023). Liang, D. D, & Zhang, Z. G. Discussion on the Causes and Prevention of Rocky Desertification in Guizhou Province. J. Anhui Agri Sci. 28, 12490–12491, doi: 10.13989/j.cnki.0517–6611.2008.28.023 (2008). Yang, R. Y., Zhong, C.B., Yang, Z.S., Liu, F.L., & Peng, H.Y. Analysis on poverty influencing factors in deep poverty county of Karst Rocky-desertified Area in Southwest China. World Regional Studies. 31(6), 1298–1309, http://dx.doi.org/10.3969/j.issn.1004–9479.2022.06.2020533 (2022). Xie, Y. C., Lin, S. Y., Tu, S. S., L, Y., & Pan, X. C. Identification and spatial pattern of multidimensional poverty measurement in karst rocky desertification regions. Trans CSAE. 36, 276–285. http://dx.doi.org/10.11975/j.issn.1002–6819.2020.22.031 (2020). Chinese Pharmacopoeia Commission. Chinese Pharmacopoeia edition Volume I . 56. (2020). Du, Y., Zhou, J., Wu, Y. F., & OuYang, Y. X. Cultivation technology of Asparagus lycopodineus : a preliminary study. Cont. Hort. 45, 62–63 + 66. doi: 10.14051/j.cnki.xdyy.2022.05.036 (2022). LEI, L. H., CHEN, Y., OU, L. J., XU, Y., & YU, X. Aqueous root extract of Asparagus cochinchinensis (Lour.) Merr. Has antioxidant activity in D-galactose-induced aging mice. BMC Complementary and Alternative Medicine. 17(1): 469. http://dx.doi.org/10.1186/s12906-017-1975-x (2017). LEE, H. A. et al . Asparagus cochinchinensis stimulates release of nerve growth factor and abrogates oxidative stress in the Tg2576 model for Alzheimer’s disease. BMC Complementary and Alternative Medicine. 18(1): 125 https://doi.org/10.1186/s12906-017-1775–3 (2018). Kim, J. Y., Choi, H. Y., Kim, H. M., Choi, J. H, & Jang, D. S. A Novel Cytotoxic Steroidal Saponin from the Roots of Asparagus cochinchinensis . Plants. 10(10), 2067. https://doi.org/10.3390/plants10102067 (2021). Liang, H. L. A. et al . Effects of Miao Medicine Asparagus on colorectal cancer cell proliferation and migration. Chin. J. Gerontol. 41(23), 5249–5252. http://dx.doi.org/10.3969/j.issn.1005–9202.2021.23.022(2021 ). Liu, M. M., Zhou, Y. Z., Yuan, L. M., Shui, J. R., Xiong, L., & Bian, M. H. Brewing and quality analysis of Asparagus new rice wine. Food Ferment. Ind. 45(1), 137–144. http://dx.doi.org/10.13995/j.cnki.11–1802/ts.018083 (2019). Zeng, G. P., Liu, H. C., & Zhang, X. Suitable Environment of Asparagus cochinchinensis in Guizhou. Guizhou Agri Sci. 38, 48–50. https://www.cabdirect.org/cabdirect/abstract/20103170869 (2010). Deng, L. F., Yuan, H. R., Xie, J., Ge, L. F, & Chen, Y. Herbaceous plants are better than woody plants for carbon sequestration. Resources. Conserv. Recycl. 184, 106431. https://doi.org/10.1016/j.resconrec.2022.106431 (2022). Gilardelli, F. et al . Root Characteristics of Herbaceous Species for Topsoil Stabilization in Restoration Projects. Land Degrad. Dev. 28(7), 2074–2085. https://doi.org/10.1002/ldr.2731 (2017). Yang, K. T., Han, X. M., & Liiang, Y. F. Effects of Planting Three Kinds of Medicinal Materials under the Forest on Soil Physicochemical Properties and Nutrients, J.Anhui Agric.Sci. 51(9): 83–86. http://dx.doi.org/10.3969/j.issn.0517–6611.2023.09.020 (2023). Liu, C. et al . Plant adapt ability in karst regions. J. Plant Res. 134, 889–906. http://dx.doi.org/10.1007/s10265-021-01330-3 (2021). Song, H. Y., Zhang, J., Zhao, Y. J., Teng, J., & Liu, J. C. Effects of rocky desertification on growth and biomass accumulation and distribution of terminal twigs in Viburnum chinshanense Graeb n. Plant Sci. J. 36(1), 103–111. http://dx.doi.org/10.11913/PSJ.2095–0837.2018.10103 (2018). Zhang, X., Liu, H. C., & Zeng, G. P. Research on the Effect of Different Water Supply on Resistance Physiology and Appearance Charactersin Asparagus Cochinchinensis (Lour.) Merr. Modern Chin. Med. 12, 19–22. http://dx.doi.org/10.13313/j.issn.1673–4890.2010.02.002 (2010). Huang, R. Y. et al . Responses of Physiology, Photosynthetic Characteristics and Leaf surface Micromorphology of Berberis poiretii to Drought Stress. Acta Agrestia Sin. 31, 1386–1392. doi: 10.11733/j.issn.1007–0435.2023.05.013 (2023). Liu, C. C., Liu, Y. G., Guo, K., Fan, Y., Li, G. Q., & Zheng, Y. R. Effect of drought on pigments, osmotic adjustment and antioxidant enzymes in six woody plant species in karst habitats of southwestern China. Environ. Exp. Bot. 71(2), 174–183. https://doi.org/10.1016/j.envexpbot.2010.11.012 (2011). Pan, R. Z., Wang, X. Q., & Li, N. H. Plant Physiology , seventh ed. 81 (Higher Education Press, 2012). Hou, R. R. et al . Polysaccharide from Echinacea purpurea reduce the oxidant stress in vitro and in vivo. Int. J. Biol. Macromol. 149, 41–50. https://doi.org/10.1016/j.ijbiomac.2020.01.129 (2020). Huang, H. L., Chen, F., Long, R., & Huang, J. L. The antioxidant activities in vivo of bitter gourd polysaccharide. J. Biol. Macromol. 145, 141–144. https://doi.org/10.1016/j.ijbiomac.2019.12.165 (2020). Lin, Y. Y. et al . Enzyme and microwave coassisted extraction, structural characterization and antioxidant activity of polysaccharides from Purple-heart Radish. Food Chemistry. 372, 131274. http://dx.doi.org/10.1016/j.foodchem.2021.131274 (2022). Wan, L. Y., Fu, J. E., Pan, L. M., Ke, F., & Wei, S. G. Analysis and evaluation of agronomic traits of Guangxi Asparagus germplasm resources. Jiangsu Agri Sci. 48, 124–128. http://dx.doi.org/10.15889/j.issn.1002–1302.2020.24.023 (2020). Peng, X., Huang, Y., Che, J. X., & Lu, L. Changing rule of soil physicochemical features under different grade of desertification A case study at Puding station of Guizhou karst center. China Karst. 28(4): 402–405. http://dx.doi.org/doi:10.3969/j.issn.1001–4810.2009.04.012 (2009). Viviana, L. P., Engil, P. P., Raphaël, W., Marcel, H., & Johan, S. Improvement of soil structure through organic crop management, conservation tillage and grass-clover ley. Soil Tillage Res. 180, 1–9. http://dx.doi.org/10.1016/j.still.2018.02.007 (2018). Manlay, R. J., Feller, C., & Swift, M. J. Historical evolution of soil organic matter concepts and their relationships with the fertility and sustainability of cropping systems. Agric, Ecosyst. Environ. 119, 217–233. https://doi.org/10.1016/j.agee.2006.07.011 (2007). Wu, Y. N., Yu. L. F., Zhang, L. M., Liu, N., & Yan, L. B. Characteristics and influencing factors of soil carbon pool during vegetation restoration in Karst Plateau. Ecol. Environ. Sci. 29, 1935–1942. https://www.jeesci.com/EN/Y2020/ V29/I10/1935 (2020). Zhang, W., Zhao, J., Pan, F., Li, D. J., & Chen, H. S. Changes in nitrogen and phosphorus limitation during secondary succession in a karst region in southwest China. Plant Soil. 391, 77–91. http://dx.doi.org/10.1007/s11104-015-2406–8 (2015). Pang, L., Wu, X., Zou, Q. L., & Li, J.X. Distribution of N and P of Common Ferns and their Relationship with Soil Supply in Qianzhong Karst Area. J Northeast For Univ. 49, 53–57. http://dx.doi.org/10.13759/j.cnki. dlxb.2021.06.011 (2021). Ming, X., & Chen, W. N. Stomata Distribution in Leaves of Different Plants. Anhui Agri Sci Bull. 27(20), 22–25. http://dx.doi.org/10.16377/j.cnki.issn1007–7731.2021.20.009 (2021). Li, H. S. Principles of plant physiological and biochemical experimental techniques . 134–137. (Higher Education Press, 2000). Li, M., Fei, Y., Li, L. X., & Fang, D. S. Method Research to Assay Total Saponin Contentin Asparagus Cochinchinese Root. J. Chengdu Univ TCM. 4, 46–48. (2001). Wu, F., Liu, M. D., Li, Y. X., & Jin, C.S. Study on Extraction Technology of Total Amino Acids and Polysaccharides from Asparagus . J Chizhou Univ. 31(3), 60–63. http://dx.doi.org/10.13420/j.cnki.jczu . 2017.03.0021 (2017). Bao D. S. Soil Agrochemical Analysis . third ed. 14–118. (Beijing Agricultural Press, 2013). Hu, Y., Wang, Y., & Li, G. Soil Nutrient Characteristics and Fertility Grade Evaluation of Newly-increased Farmland in Weibei Dryland: A Case Study of Heyang County. Chin. Agri. Sci Bull. 38, 94–100. doi: 10.11924/j.issn.1000–6850.casb2021–0886 (2022). Additional Declarations No competing interests reported. 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-4318709","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":296948884,"identity":"20eb8f1b-580a-41a9-8512-78508559c8f4","order_by":0,"name":"Mingsheng Zhang","email":"","orcid":"","institution":"Guizhou University","correspondingAuthor":false,"prefix":"","firstName":"Mingsheng","middleName":"","lastName":"Zhang","suffix":""},{"id":296948886,"identity":"6cac8da7-73ed-4bf9-a271-e9b130183105","order_by":1,"name":"liu tang","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA10lEQVRIiWNgGAWjYLCCBwYMcmzszQcOfPhBhGoeEJFgwGDMx3Ms8eDMHqK1MDAkzpPIMT7MwUaEFnv2s4dfJBQcTmyTyPlwGGiCPL/YAQK28OSlWSQYHDZu43m74XCBBYPhzNkJhByWY2YA1CLbxp674fAMHqC/bhPSwv8GrIWxjSHnwWEeNmK0AH39AKhFsY0jh4FILTfemAGVpRuz8RwzAAayBGG/sPfnGH/48MdaTr69+fGHDz9s5PmlCWgBAjYJJI4ETmXIgPkDUcpGwSgYBaNg5AIATeBE/cvXaRoAAAAASUVORK5CYII=","orcid":"","institution":"Guizhou University","correspondingAuthor":true,"prefix":"","firstName":"liu","middleName":"","lastName":"tang","suffix":""},{"id":296948888,"identity":"c79d9a62-dc47-4ef0-829d-8a9d11210c2c","order_by":2,"name":"miao liu","email":"","orcid":"","institution":"Guizhou University","correspondingAuthor":false,"prefix":"","firstName":"miao","middleName":"","lastName":"liu","suffix":""},{"id":296948889,"identity":"e6bb6e3e-9e0c-45f5-bc01-304c3d05d904","order_by":3,"name":"liyang shangguan","email":"","orcid":"","institution":"Guizhou University","correspondingAuthor":false,"prefix":"","firstName":"liyang","middleName":"","lastName":"shangguan","suffix":""},{"id":296948890,"identity":"1fde2d74-13fc-475e-b49c-f42100197fbf","order_by":4,"name":"haidong Huang","email":"","orcid":"","institution":"Guizhou University","correspondingAuthor":false,"prefix":"","firstName":"haidong","middleName":"","lastName":"Huang","suffix":""},{"id":296948892,"identity":"606c3186-8241-4c5a-824e-13100574d012","order_by":5,"name":"gonggu lv","email":"","orcid":"","institution":"Guizhou University","correspondingAuthor":false,"prefix":"","firstName":"gonggu","middleName":"","lastName":"lv","suffix":""},{"id":296948894,"identity":"61b30a37-4f4d-4852-b915-75eca34f0709","order_by":6,"name":"Ye hang","email":"","orcid":"","institution":"Guizhou University","correspondingAuthor":false,"prefix":"","firstName":"Ye","middleName":"","lastName":"hang","suffix":""}],"badges":[],"createdAt":"2024-04-24 13:53:29","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-4318709/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4318709/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":65808103,"identity":"e961d852-b9f5-4513-9223-96b69a44e297","added_by":"auto","created_at":"2024-10-03 03:38:46","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":753275,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4318709/v1/fb7e0f4f-063c-4e2b-93fd-1ae241915ef5.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Adaptation of Asparagus munitus to karst adversity and its effects on soil","fulltext":[{"header":"Introduction","content":"\u003cp\u003eAccording to the latest karst rocky desertification (KRD) survey, China's KRD area was 7.22\u0026nbsp;million hm2 in 2021\u003csup\u003e[\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e]\u003c/sup\u003e. KRD severely limits human land use due to poor nutrient storage, severe temporary drought, and poor ecological stability \u003csup\u003e[\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e]\u003c/sup\u003e. Studies have shown a correlation between poverty and the KRD environment \u003csup\u003e[\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e, \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e]\u003c/sup\u003e. To address ecological and economic issues, controlling KRD is urgently needed for environmental protection and poverty eradication.\u003c/p\u003e \u003cp\u003e \u003cem\u003eAsparagus cochinchinensis\u003c/em\u003e (Lour.) Merr. is a perennial herb; its tuberous roots have long been used in traditional Chinese medicine and was included in the 2020 Chinese Pharmacopoeia \u003csup\u003e[\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e]\u003c/sup\u003e. Currently, \u003cem\u003eAsparagus munitus\u003c/em\u003e Wang et S.C. Chen (\u003cem\u003eA. munitus\u003c/em\u003e), \u003cem\u003eAsparagus lycopodineus\u003c/em\u003e (Baker.) Wang et. Tang (\u003cem\u003eA. lycopodineus\u003c/em\u003e) and \u003cem\u003eAsparagus subscandens\u003c/em\u003e Wang et S.C. Chen (\u003cem\u003eA. subscandens\u003c/em\u003e) have been artificially domesticated \u003csup\u003e[\u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e6\u003c/span\u003e]\u003c/sup\u003e. Although only \u003cem\u003eA. cochinchinensis\u003c/em\u003e is officially recognized, the other three \u003cem\u003eAsparagus\u003c/em\u003e species are more widely cultivated due to \u003cem\u003eA. cochinchinensis\u003c/em\u003e's low yield and their similar medicinal composition.\u003c/p\u003e \u003cp\u003eWith Asparagus development and utilization, there are great market prospects in pharmacology \u003csup\u003e[\u003cspan additionalcitationids=\"CR8 CR9\" citationid=\"CR7\" class=\"CitationRef\"\u003e7\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e]\u003c/sup\u003e and health foods \u003csup\u003e[\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e]\u003c/sup\u003e. \u003cem\u003eA. munitus\u003c/em\u003e's wide distribution indicates strong environmental adaptability \u003csup\u003e[\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e]\u003c/sup\u003e. Several studies have shown that herbaceous plants undergo better carbon sequestration than woody plants \u003csup\u003e[\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e]\u003c/sup\u003e, and their well-developed root systems are able to fix the topsoil layer and prevent soil erosion \u003csup\u003e[\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e]\u003c/sup\u003e. The effects of planting \u003cem\u003eAsparagus\u003c/em\u003e on soil physicochemical properties and nutrients have been studied \u003csup\u003e[\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e]\u003c/sup\u003e, but few studies exist on \u003cem\u003eAsparagus\u003c/em\u003e planting in karst soil and its interaction.\u003c/p\u003e \u003cp\u003eThis study compared \u003cem\u003eA. munitus\u003c/em\u003e changes in KRD and arable land (AL) to predict survival strategies in KRD. It also tested whether \u003cem\u003eA. munitus\u003c/em\u003e could improve KRD soil conditions, providing a basis for its cultivation and KRD control and new ideas for medicinal plant development.\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003e \u003cb\u003eA. munitus\u003c/b\u003e \u003cb\u003eunder KRD conditions shows a decreasing trend\u003c/b\u003e\u003c/p\u003e \u003cp\u003eThe results revealed significant differences (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05) in stem number, stem length, middle lateral branch length and leaf length between the EG and CK plants of \u003cem\u003eA. munitus\u003c/em\u003e (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). No significant differences were observed for the other indicators. LSD indicated no significant difference in below-ground biomass, however CK averaged approximately one-third higher than EG, EG tubers tended to be shorter but greater in diameter (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e2\u003c/span\u003eA, B). Overall, EG growth was inhibited, especially above ground, and was weaker than that in the CK treatment.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eAgronomic traits of \u003cem\u003eA. munitus\u003c/em\u003e plants grown in different regionsa and b indicate significant differences at the 0.05 level (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05); A and B indicate significant differences at the 0.01 level (p\u0026thinsp;\u0026lt;\u0026thinsp;0.01). The values are the means\u0026thinsp;\u0026plusmn;\u0026thinsp;standard deviations\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"4\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTrait\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eUnit\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eControl check\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eExperiment group\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eNumber of stems\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eeach\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e5.33\u0026thinsp;\u0026plusmn;\u0026thinsp;0.82 A\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3.67\u0026thinsp;\u0026plusmn;\u0026thinsp;0.52 B\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBasal stem thickness\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003emm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3.05\u0026thinsp;\u0026plusmn;\u0026thinsp;0.46 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.98\u0026thinsp;\u0026plusmn;\u0026thinsp;0.53 a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eStems length\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ecm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e141.66\u0026thinsp;\u0026plusmn;\u0026thinsp;16.8 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e114.05\u0026thinsp;\u0026plusmn;\u0026thinsp;21.0 b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLateral branch length\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ecm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e21.97\u0026thinsp;\u0026plusmn;\u0026thinsp;2.84 A\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e16.85\u0026thinsp;\u0026plusmn;\u0026thinsp;1.88 B\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLeafy shoot length\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003emm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e6.76\u0026thinsp;\u0026plusmn;\u0026thinsp;0.87 A\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e5.18\u0026thinsp;\u0026plusmn;\u0026thinsp;0.61 B\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAbove-ground biomass\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eg\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e42.30\u0026thinsp;\u0026plusmn;\u0026thinsp;10.19 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e34.88\u0026thinsp;\u0026plusmn;\u0026thinsp;6.64 a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eUnderground biomass\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eg\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e245.53\u0026thinsp;\u0026plusmn;\u0026thinsp;62.67 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e186.76\u0026thinsp;\u0026plusmn;\u0026thinsp;30.11 a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cb\u003eBiochemical responses of\u003c/b\u003e \u003cb\u003eA. munitus\u003c/b\u003e \u003cb\u003eleafy shoots to KRD\u003c/b\u003e\u003c/p\u003e \u003cp\u003eThe stomatal density in the EG treatment was significantly greater than that in the CK treatment (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e3\u003c/span\u003e), averaging 471.26 mm\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e. Compared to those in the CK treatment, the photosynthetic pigment content in the EG treatment was markedly lower; however, the carotenoid-to-chlorophyll ratio differed highly significantly between the groups (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Similarly, the leaf SOD, POD, and CAT activities in the EG treatment were 1.4, 1.6 and 1.7 times greater than those in the CK treatment, respectively. Osmoregulatory substances showed similar increasing trends in the EG compared to the CK. These results suggest that \u003cem\u003eA. munitus\u003c/em\u003e individuals experience some degree of stress under KRD conditions.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eBiochemical indicators of plants in different environments. A and B indicate significant differences at the 0.01 level (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.01). The values are the means\u0026thinsp;\u0026plusmn;\u0026thinsp;standard deviations. FW: fresh weight\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"4\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTrait\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eUnit\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eControl check\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eExperiment group\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eStomata density\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePcs/mm\u003csup\u003e2\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e375.43\u0026thinsp;\u0026plusmn;\u0026thinsp;45.92 B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e471.26\u0026thinsp;\u0026plusmn;\u0026thinsp;103.52 A\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eChlorophyll a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003emg/g FW\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.81\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03 A\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.43\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04 B\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eChlorophyll b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003emg/g FW\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.72\u0026thinsp;\u0026plusmn;\u0026thinsp;0.058 A\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.52\u0026thinsp;\u0026plusmn;\u0026thinsp;0.033 B\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCarotenoid\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003emg/g FW\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.41\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002 A\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.37\u0026thinsp;\u0026plusmn;\u0026thinsp;0.004 B\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCarotenoids/chlorophyll\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e16.27\u0026thinsp;\u0026plusmn;\u0026thinsp;0.12 B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e18.83\u0026thinsp;\u0026plusmn;\u0026thinsp;0.49 A\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cb\u003eThe medicinal components of\u003c/b\u003e \u003cb\u003eA. munitus\u003c/b\u003e \u003cb\u003eare not affected by KRD\u003c/b\u003e\u003c/p\u003e \u003cp\u003eTotal saponins, polysaccharides and total amino acids are the main medicinal components of \u003cem\u003eA. munitus\u003c/em\u003e tubers. The results showed that the polysaccharide content was significantly greater (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05) in the EG tubers than in the CK tubers, while no significant differences were observed for the other components (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e4\u003c/span\u003e). These findings indicate that \u003cem\u003eA. munitus\u003c/em\u003e plants can accumulate medicinal components normally when planted in KRD areas, which is even slightly higher than what is observed in the CK treatment.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cb\u003eCorrelation analysis and principal component analysis of agronomic traits in\u003c/b\u003e \u003cb\u003eA. munitus\u003c/b\u003e Correlation analysis revealed strong associations between most agronomic traits, except for basal stem thickness, which was less correlated with the other indicators (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e5\u003c/span\u003e). After removing the basal stem thickness, the remaining traits were subjected to principal component analysis. The first two principal components had eigenvalues above 1 (5.248 and 1.424) and accounted for 83.400% of the total variance (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Thus, these two components represent the majority of the information from the eight agronomic traits. Leafy shoot length and tuber diameter had the highest eigenvector values for the first and second principal components (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). Therefore, the principal components included leafy shoot length and tuber diameter.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003ePrincipal component analysis of the agronomic traits of \u003cem\u003eA. munitus\u003c/em\u003e\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"4\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eComponents\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eEigenvalue\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eContribution rate\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eCumulative contribution rate\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e5.248\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e65.594\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e65.594\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1.424\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e17.806\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e83.400\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0.513\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e6.411\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e89.811\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0.343\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e4.289\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e94.100\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0.258\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e3.221\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e97.320\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0.119\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1.485\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e98.806\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0.077\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.968\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e99.773\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0.018\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.227\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e100.000\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab4\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 4\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003ePrincipal component analysis eigenvectors and contributions of agronomic traits\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"3\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eAgronomic trait\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e \u003cp\u003eEigenvector\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePrincipal Component 1\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePrincipal Component 2\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eNumber of stems\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0.867\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e-0.198\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eStems length\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0.782\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.321\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLateral branch length\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0.851\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.160\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLeafy shoot length\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0.976\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e-0.010\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAbove-ground biomass\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0.793\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.540\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eUnderground biomass\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0.951\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.083\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTuber length\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0.821\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e-0.228\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTuber diameter\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e-0.025\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.951\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab5\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 5\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eEffect of \u003cem\u003eA. munitus\u003c/em\u003e planting on the physical properties of KRD soils. a and b indicate significant differences at the 0.05 level (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05); A and B indicate significant differences at the 0.01 level (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.01). The values are the means\u0026thinsp;\u0026plusmn;\u0026thinsp;standard\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"4\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTrait\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eUnit\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSoil control\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSoil near \u003cem\u003eA. munitus\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003epH\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e7.37\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05 A\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e7.21\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06 B\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003esoil water content\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e24.50\u0026thinsp;\u0026plusmn;\u0026thinsp;1.10 B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e30.76\u0026thinsp;\u0026plusmn;\u0026thinsp;0.97 A\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003esoil bulk density\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.93\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02 b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.02\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04 a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003esoil total porosity\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e59.81\u0026thinsp;\u0026plusmn;\u0026thinsp;4.24 B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e62.15\u0026thinsp;\u0026plusmn;\u0026thinsp;0.92 A\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003esoil capillary porosity\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e54.83\u0026thinsp;\u0026plusmn;\u0026thinsp;1.48 b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e58.78\u0026thinsp;\u0026plusmn;\u0026thinsp;0.89 a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003esoil noncapillary porosity\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.64\u0026thinsp;\u0026plusmn;\u0026thinsp;0.21 B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3.37\u0026thinsp;\u0026plusmn;\u0026thinsp;0.12 A\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab6\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 6\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eEffect of \u003cem\u003eA. munitus\u003c/em\u003e planting on the soil nutrient content of KRD soils. a and b indicate significant differences at the 0.05 level (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05), and A and B indicate significant differences at the 0.01 level (\u003cem\u003ep\u0026thinsp;\u0026lt;\u003c/em\u003e\u0026thinsp;0.01). The values are the means\u0026thinsp;\u0026plusmn;\u0026thinsp;standard deviations\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"4\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTrait\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eUnit\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSoil control\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSoil near A. munitus\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSoil organic matter\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e5.80\u0026thinsp;\u0026plusmn;\u0026thinsp;0.23 B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e8.09\u0026thinsp;\u0026plusmn;\u0026thinsp;0.40 A\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTotal nitrogen\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eg/kg\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3.15\u0026thinsp;\u0026plusmn;\u0026thinsp;0.17 B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e4.04\u0026thinsp;\u0026plusmn;\u0026thinsp;0.21 A\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTotal phosphorus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eg/kg\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.85\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.72\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05 b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTotal potassium\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eg/kg\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e12.82\u0026thinsp;\u0026plusmn;\u0026thinsp;0.12 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e11.07\u0026thinsp;\u0026plusmn;\u0026thinsp;0.54 b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAlkali-hydro nitrogen\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eg/kg\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e243.87\u0026thinsp;\u0026plusmn;\u0026thinsp;10.04 B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e317.72\u0026thinsp;\u0026plusmn;\u0026thinsp;13.14 A\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAvailable phosphorus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003emg/kg\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e6.26\u0026thinsp;\u0026plusmn;\u0026thinsp;0.75 A\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.32\u0026thinsp;\u0026plusmn;\u0026thinsp;0.19 B\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAvailable potassium\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003emg/kg\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e119.33\u0026thinsp;\u0026plusmn;\u0026thinsp;10.02 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e94.33\u0026thinsp;\u0026plusmn;\u0026thinsp;6.43 b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cb\u003eA. munitus\u003c/b\u003e \u003cb\u003ePlanting Changes the Soil Physical Properties in the KRD Area\u003c/b\u003e\u003c/p\u003e \u003cp\u003eThe KRD test site was weakly alkaline. Planting \u003cem\u003eA. munitus\u003c/em\u003e significantly reduced the SA pH by 0.16 units on average compared to that in SC (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.01). Additionally, SA had an extremely significantly higher SWC versus SC (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.01), at more than 31%. SBD, SP, SCP and SNP all differed significantly between SA and SC to varying degrees. These results demonstrated that \u003cem\u003eA. munitus\u003c/em\u003e strongly affects the basic physical properties of KRD soils.\u003c/p\u003e \u003cp\u003e \u003cb\u003ePlanting\u003c/b\u003e \u003cb\u003eA. munitus\u003c/b\u003e \u003cb\u003ealtered the KRD soil nutrient content\u003c/b\u003e\u003c/p\u003e \u003cp\u003eThe KRD soil nutrient levels varied according to fertility grade. SOM and TN were enriched (Class I), AK was moderate (Class III), and AP was depleted (Class IV), indicating that the site was relatively deficient in P and K. Planting \u003cem\u003eA. munitus\u003c/em\u003e significantly increased SOM, TN and AN in SA versus SC (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05). TP and AP differed significantly between SA and SC, with SC having 2.7 times greater AP. TK and AK were also significantly greater in the SC treatment than in the SA treatment (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05). Taken together, these findings demonstrated that \u003cem\u003eA. munitus\u003c/em\u003e planting depletes soil P and K, especially P. Overall, \u003cem\u003eA. munitus\u003c/em\u003e decreased the KRD soil pH but increased the SWC, SBD, SP, SOM, TN and AN while decreasing P and K. These data indicate that planting \u003cem\u003eA. munitus\u003c/em\u003e significantly alters basic physicochemical properties of KRD soil, largely in beneficial ways.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003e \u003cb\u003eAdaptation of\u003c/b\u003e \u003cb\u003eA. munitus\u003c/b\u003e \u003cb\u003eto KRD habitats\u003c/b\u003e\u003c/p\u003e \u003cp\u003eTo adapt to the harsh conditions of KRD habitats, plants exhibit a range of altered morphological and physiological characteristics \u003csup\u003e[\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e]\u003c/sup\u003e. \u003cem\u003eA. munitus\u003c/em\u003e exhibited shortened stems, branches, and leaves and a reduced aboveground biomass in KRD areas, indicating an overall \"shrinkage\" phenotype. This finding aligns with that of other KRD-adapted plant species \u003csup\u003e[\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e]\u003c/sup\u003e and may be linked to the seasonal drought commonly experienced in these areas \u003csup\u003e[\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e]\u003c/sup\u003e. The significantly greater stomatal density observed in the EG mirrors similar adaptive increases under drought conditions reported in \u003cem\u003eBerberis poiretii\u003c/em\u003e \u003csup\u003e[\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e]\u003c/sup\u003e. By reducing the number of leafy shoots and the aboveground light-exposed area while increasing stomatal transpiration per unit area, \u003cem\u003eA. munitus\u003c/em\u003e may be minimizing water loss and heat damage. This shrinkage strategy could aid survival in the water-limited KRD habitats. Compared to those in the CK treatment, plants in the EG treatment had significantly lower total chlorophyll but a greater carotenoid-to-chlorophyll ratio. This finding aligns with that of other karst species under drought stress \u003csup\u003e[\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e]\u003c/sup\u003e and may be attributed to the photoprotective role of carotenoids in addition to their functions in light absorption and energy transfer \u003csup\u003e[\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e21\u003c/span\u003e]\u003c/sup\u003e. Such adaptations enable the plants to thrive in the challenging KRD environment.\u003c/p\u003e \u003cp\u003eThe results of the antioxidant enzyme activity and osmoregulatory substance content analyses showed that plants were subjected to a certain degree of adverse stress in the KRD habitat \u003csup\u003e[\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e]\u003c/sup\u003e. Tests of the medicinal properties of the harvested tubers showed that the polysaccharide content in the EG treatment was significantly higher than that in the CK treatment, which could be related to the physiological functions of polysaccharides \u003csup\u003e[\u003cspan additionalcitationids=\"CR23\" citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e]\u003c/sup\u003e. Correlation analysis and principal component analysis of the agronomic traits of \u003cem\u003eA. munitus\u003c/em\u003e revealed strong correlations among most agronomic indicators except for basal stems. These results could be simplified into two principal components, leafy shoot length and tuber diameter, which was similar to the results of a previous study \u003csup\u003e[\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e]\u003c/sup\u003e, suggesting that the belowground portion of \u003cem\u003eA. munitus\u003c/em\u003e may play a more vital role in the plant's growth and development.\u003c/p\u003e\n\u003ch3\u003ePlanting A. munitus to improve KRD soil conditions\u003c/h3\u003e\n\u003cp\u003eThe results of the study revealed that the planting of \u003cem\u003eA. munitus\u003c/em\u003e significantly reduced the soil pH and augmented the SWC, providing a favorable environment for plant survival. However, the SBD was greater in the SA treatment group than in the control group, unlike previous findings \u003csup\u003e[\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e]\u003c/sup\u003e. This may be due to the high gravel and stone contents at the sampling site affecting the results. SP, SCP and SNP were significantly greater in SA than in SC, which was potentially related to root growth or increased SOM improving soil structure and looseness \u003csup\u003e[\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e]\u003c/sup\u003e. SOM, TN and AN were significantly higher in SA than in SC. SOM is central to soil fertility and vegetation recovery \u003csup\u003e[\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e, \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e]\u003c/sup\u003e. Related studies have shown that N limits early-mid-stage karst ecosystem recovery \u003csup\u003e[\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e]\u003c/sup\u003e. Fern rhizosphere soils have lower AP than non-rhizosphere soils \u003csup\u003e[\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e]\u003c/sup\u003e, but the difference here was greater. This could be because of the already low substrate P concentration coupled with the high P demand during \u003cem\u003eA. munitus\u003c/em\u003e growth. Similarly, low soil P in \u003cem\u003eA. munitus\u003c/em\u003e habitats was previously observed \u003csup\u003e[\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e]\u003c/sup\u003e, suggesting that P depletion is linked to high P requirements and needs further verification. Overall, \u003cem\u003eA. munitus\u003c/em\u003e decreased the KRD soil pH but increased the SWC, SBD, SP, SOM, TN and AN while decreasing P and K. The data indicate that planting \u003cem\u003eA. munitus\u003c/em\u003e significantly alters the basic physical and nutritional properties of KRD soil, largely in beneficial ways.\u003c/p\u003e \u003cp\u003eThe results of the study revealed that the planting of \u003cem\u003eA. munitus\u003c/em\u003e significantly reduced the soil pH and augmented the SWC, providing a favorable environment for plant survival. However, an unexpected outcome was noted in the SBD, which was more pronounced in the treatment group with \u003cem\u003eA. munitus\u003c/em\u003e (SA) than in the control group, deviating from the findings of Peng et al \u003csup\u003e[\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e]\u003c/sup\u003e. This anomaly could be attributed to the high concentration of gravel and stones at the sampling site, which might have influenced the results. The SP, SCP, and SNP were markedly higher in SA than in the control group, which was potentially associated with enhanced root growth or improved soil structure and looseness as per Viviana et al \u003csup\u003e[\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e]\u003c/sup\u003e. The SOM, TN, and AN were significantly elevated in SA compared to SC. SOM is pivotal for soil fertility \u003csup\u003e[\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e]\u003c/sup\u003e and vegetation recovery \u003csup\u003e[\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e]\u003c/sup\u003e. Related studies have revealed that nitrogen constraints early-mid-stage karst ecosystem recovery \u003csup\u003e[\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e]\u003c/sup\u003e. Fern rhizosphere soils exhibit lower AP than non-rhizosphere soils \u003csup\u003e[\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e]\u003c/sup\u003e, but the difference observed in this study was more pronounced. This could be attributed to the already low substrate P concentration coupled with the high P demand during \u003cem\u003eA. munitus\u003c/em\u003e growth. Similarly, low soil P in \u003cem\u003eA. munitus\u003c/em\u003e habitats was previously observed \u003csup\u003e[\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e]\u003c/sup\u003e, suggesting that P depletion is linked to high P requirements and necessitates further verification. In summary, \u003cem\u003eA. munitus\u003c/em\u003e reduced the KRD soil pH but increased the SWC, SBD, SP, SOM, TN, and AN while decreasing P and K. The data underscore that planting \u003cem\u003eA. munitus\u003c/em\u003e significantly alters the basic physical and nutritional properties of KRD soil, predominantly in beneficial ways.\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eIn conclusion, this experiment initially confirmed that \u003cem\u003eA. munitu\u003c/em\u003e could thrive in the KRD environment. This was evident from the morphological changes and physio-chemical reactions that occurred in the plant and its surrounding environment. After planting \u003cem\u003eA. munitu\u003c/em\u003e, the physical and chemical properties of KRD soil changed, with increased soil water holding capacity and overall fertility, making it more suitable for other plants. Planting \u003cem\u003eA. munitu\u003c/em\u003e in KRD areas not only brings economic benefits but also improves KRD soil conditions and ecological value. This lays a good foundation for KRD management and sustainable development in karst areas.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cdiv id=\"Sec7\" class=\"Section2\"\u003e\n \u003ch2\u003eTest site and plant materials\u003c/h2\u003e\n \u003cp\u003eThe test area is located at the experimental base (26\u0026deg;25\u0026prime;54\u0026Prime;N, 106\u0026deg;40\u0026prime;53\u0026Prime;E) of Guizhou University, Huaxi District, Guiyang City, Guizhou Province, China. This area features a karst plateau landform with a humid subtropical monsoon climate, an average elevation of 1178 m, an average annual temperature of 20.21\u0026deg;C, and an average annual precipitation of 1,318.90 mm. The vegetation is mainly natural secondary forest. The germplasm resources of \u003cem\u003eA. munitus\u003c/em\u003e were sourced from the Planting Base of \u003cem\u003eAsparagus\u003c/em\u003e in Xingren Town of Qiandongnan (in Guizhou Province, China), and were identified by Prof. Zhang Mingsheng of Guizhou University. The plants were sampled on April 20, 2023 (Fig. \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eThe effect of karst environments on\u003c/strong\u003e \u003cstrong\u003eA. Munitus.\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eMeasurement of agronomic traits\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eA. munitus\u003c/em\u003e plants were randomly selected from the KRD habitat as the experimental group (EG), and plants from the AL habitat as the control (CK). The number of plant stems, basal stem thickness (mm), stem length (cm), lateral branch length (cm), foliage branch length (mm), tuber length (cm) and diameter (cm) were measured. an electronic balance was used for biomass measurements (the dry parts and sediment attached to the plants were removed, and the plants were subsequently dried in an electric oven at 60\u0026deg;C until the weight was constant).\u003c/p\u003e\n\u003c/div\u003e\n\u003ch3\u003eMeasurement of physio-biochemical indicators\u003c/h3\u003e\n\u003cp\u003eThe leafy shoots of \u003cem\u003eA. munitus\u003c/em\u003e were randomly collected and rapidly frozen in liquid nitrogen to the laboratory. The stomata density was calculated according to the methods of Ming and Chen \u003csup\u003e[\u003cspan class=\"CitationRef\"\u003e32\u003c/span\u003e]\u003c/sup\u003e. The chlorophyll content in the leafy shoots was measured according to the methods of Li \u003csup\u003e[\u003cspan class=\"CitationRef\"\u003e33\u003c/span\u003e]\u003c/sup\u003e. Superoxide dismutase (SOD), peroxidase (POD), catalase (CAT), soluble sugar, and proline were determined by using a kit (Suzhou Keming Biotechnology Co., Ltd., China), and the soluble protein content was determined by the Caulmers Brilliant Blue method with reference to Li \u003csup\u003e[\u003cspan class=\"CitationRef\"\u003e33\u003c/span\u003e]\u003c/sup\u003e.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eMeasurement of medicinal components\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eA. munitus\u003c/em\u003e tubers were collected randomly from two habitats, boiled in boiling water for 10 min, peeled, and subsequently dried at 60\u0026deg;C and crushed. The total saponin, total amino acid and polysaccharide contents of the tubers were determined by the methods of Li et al. \u003csup\u003e[\u003cspan class=\"CitationRef\"\u003e34\u003c/span\u003e]\u003c/sup\u003e and Wu Fei et al \u003csup\u003e[\u003cspan class=\"CitationRef\"\u003e35\u003c/span\u003e]\u003c/sup\u003e.\u003c/p\u003e\n\u003cdiv id=\"Sec9\" class=\"Section2\"\u003e\n \u003ch2\u003eMeasurement of soil physical properties\u003c/h2\u003e\n \u003cp\u003eOn several consecutive sunny afternoons, six soil samples near \u003cem\u003eA. munitus\u003c/em\u003e plants (SA) in the KRD area and unplanted blank controls (SC) were collected to test physicochemical properties using methods in \u0026quot;Soil Agrochemical Analysis\u0026quot; \u003csup\u003e[\u003cspan class=\"CitationRef\"\u003e36\u003c/span\u003e]\u003c/sup\u003e. Soil pH was determined by electrode method. SA and SC soils from 0\u0026ndash;20 cm were collected by ring knife method, and assessed for soil bulk density (SBD), soil water content (SWC), soil total porosity (SP), soil capillary porosity (SCP), and soil non-capillary porosity (SNP) using the following formulae: \u003cem\u003eM\u003c/em\u003e (ring knife mass), \u003cem\u003eM\u003c/em\u003e\u003csub\u003e\u003cem\u003e1\u003c/em\u003e\u003c/sub\u003e (soil fresh mass), \u003cem\u003eM\u003c/em\u003e\u003csub\u003e\u003cem\u003e2\u003c/em\u003e\u003c/sub\u003e (soil saturated mass), \u003cem\u003eM\u003c/em\u003e\u003csub\u003e\u003cem\u003e3\u003c/em\u003e\u003c/sub\u003e (soil resting mass for 12 h), \u003cem\u003eM\u003c/em\u003e\u003csub\u003e\u003cem\u003e4\u003c/em\u003e\u003c/sub\u003e (soil air-dried mass), and \u003cem\u003eV\u003c/em\u003e (ring knife volume).\u003c/p\u003e\n \u003cp\u003e\u003cspan class=\"InlineEquation\"\u003e\u003cimg 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\"\u003e\u0026nbsp;\u003c/span\u003e\u003c/p\u003e\n\u003c/div\u003e\n\u003cdiv id=\"Sec10\" class=\"Section2\"\u003e\n \u003ch2\u003eMeasurement of soil nutrients\u003c/h2\u003e\n \u003cp\u003eSoil organic matter (SOM) content was determined via potassium dichromate oxidation, total nitrogen (TN) via Kjeldahl method, total phosphorus (TP) via HClO\u003csub\u003e4\u003c/sub\u003e-H2SO\u003csub\u003e4\u003c/sub\u003e, total potassium (TK) via sodium hydroxide fusion and flame photometry, alkali-hydro nitrogen (AN) via alkaline diffusion, available phosphorus (AP) via sodium bicarbonate extraction, and molybdenum antimony anti-colorimetry, and available potassium (AK) via flame photometry method \u003csup\u003e[\u003cspan class=\"CitationRef\"\u003e36\u003c/span\u003e]\u003c/sup\u003e. The level of soil nutrient grade was evaluated according to the method of Hu et al \u003csup\u003e[\u003cspan class=\"CitationRef\"\u003e37\u003c/span\u003e]\u003c/sup\u003e.\u003c/p\u003e\n\u003c/div\u003e\n\u003cdiv id=\"Sec14\" class=\"Section2\"\u003e\u003cbr\u003e\u003c/div\u003e"},{"header":"Declarations","content":"\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\n\u003cp\u003eMS Z did supervision. LT and ML designed and conducted the experiment. LT, L S, and Y H performed collection of data and laboratory analysis. LT and M L performed statistical analysis of the data. LT, GG L and HD H were involved in writing\u0026ndash;\u0026ndash;review and editing of manuscript. All authors have read and approved the final version of manuscript.\u003c/p\u003e\n\u003ch2\u003eData Availability\u003c/h2\u003e\n\u003cp\u003eThe datasets used and/or analyzed during the current study are available from the Liu Tang on reasonable request.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCollection license statement for Cremastra appendiculata\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eStatement:\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe plant materials used in this study have been obtained with the necessary permits or licenses, complying with all applicable laws and regulations. The plant materials have been formally identified by Professor Mingsheng Zhang at Guizhou University and are deposited in the specimen repository at Guizhou University for reference and verification.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eStatement of Compliance with Endangered Species Research Policies\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe hereby affirm our commitment to adhere to relevant institutional, national, and international guidelines and legislation when conducting experimental and field research on wild plants. Specifically, we pledge to comply with the IUCN Policy Statement on Research Involving Species at Risk of Extinction and the Convention on the Trade in Endangered Species of Wild Fauna and Flora.\u003c/p\u003e\n\u003cp\u003eWe will respect principles of plant material collection and ensure strict adherence to ethical standards and best practices in the collection, use, and handling of plant samples. We will endeavor to minimize our impact on plant ecosystems in our research and mitigate any potential negative effects on endangered species.\u003c/p\u003e\n\u003cp\u003eWe will collaborate with local and international conservation organizations to ensure that our research does not result in overexploitation, misuse, or destruction of plant species. We will promote the conservation and sustainable use of plant biodiversity to support ecosystem health and human well-being.\u003c/p\u003e\n\u003cp\u003eWe solemnly pledge that our research will uphold the highest scientific standards and ethical principles to contribute positively to the protection of natural environments and biodiversity.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eDeclaration of interests\u003c/strong\u003e\u003cbr /\u003e \u0026nbsp;\u003cbr /\u003e ☑ The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.\u003cbr /\u003e \u0026nbsp;\u003cbr /\u003e ☐ The authors declare the following financial interests/personal relationships which may be considered as potential competing interests:\u003cbr /\u003e \u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eNing, X. B. et al. Main Results of the Fourth Rocky Desertification Survey in Karst Area and Analysis of Rocky Desertification Control Situation. For. Resour. Manage 3, 09\u0026ndash;14, \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://dx.doi.org/10.13466/j.cnki.lyzygl.2023.03.002\u003c/span\u003e\u003cspan address=\"10.13466/j.cnki.lyzygl.2023.03.002\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2023).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLiang, D. D, \u0026amp; Zhang, Z. G. Discussion on the Causes and Prevention of Rocky Desertification in Guizhou Province. J. Anhui Agri Sci. 28, 12490\u0026ndash;12491, doi: \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.13989/j.cnki.0517\u0026ndash;6611.2008.28.023\u003c/span\u003e\u003cspan address=\"10.13989/j.cnki.0517\u0026ndash;6611.2008.28.023\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2008).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eYang, R. Y., Zhong, C.B., Yang, Z.S., Liu, F.L., \u0026amp; Peng, H.Y. Analysis on poverty influencing factors in deep poverty county of Karst Rocky-desertified Area in Southwest China. World Regional Studies. 31(6), 1298\u0026ndash;1309, \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://dx.doi.org/10.3969/j.issn.1004\u0026ndash;9479.2022.06.2020533\u003c/span\u003e\u003cspan address=\"10.3969/j.issn.1004\u0026ndash;9479.2022.06.2020533\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2022).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eXie, Y. C., Lin, S. Y., Tu, S. S., L, Y., \u0026amp; Pan, X. C. Identification and spatial pattern of multidimensional poverty measurement in karst rocky desertification regions. Trans CSAE. 36, 276\u0026ndash;285. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://dx.doi.org/10.11975/j.issn.1002\u0026ndash;6819.2020.22.031\u003c/span\u003e\u003cspan address=\"10.11975/j.issn.1002\u0026ndash;6819.2020.22.031\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2020).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eChinese Pharmacopoeia Commission. \u003cem\u003eChinese Pharmacopoeia edition Volume I\u003c/em\u003e. 56. (2020).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDu, Y., Zhou, J., Wu, Y. F., \u0026amp; OuYang, Y. X. Cultivation technology of \u003cem\u003eAsparagus lycopodineus\u003c/em\u003e: a preliminary study. Cont. Hort. 45, 62\u0026ndash;63\u0026thinsp;+\u0026thinsp;66. doi:\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.14051/j.cnki.xdyy.2022.05.036\u003c/span\u003e\u003cspan address=\"10.14051/j.cnki.xdyy.2022.05.036\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2022).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLEI, L. H., CHEN, Y., OU, L. J., XU, Y., \u0026amp; YU, X. Aqueous root extract of \u003cem\u003eAsparagus cochinchinensis\u003c/em\u003e (Lour.) Merr. Has antioxidant activity in D-galactose-induced aging mice. BMC Complementary and Alternative Medicine. 17(1): 469. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://dx.doi.org/10.1186/s12906-017-1975-x\u003c/span\u003e\u003cspan address=\"10.1186/s12906-017-1975-x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2017).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLEE, H. A. \u003cem\u003eet al\u003c/em\u003e. \u003cem\u003eAsparagus cochinchinensis\u003c/em\u003e stimulates release of nerve growth factor and abrogates oxidative stress in the Tg2576 model for Alzheimer\u0026rsquo;s disease. BMC Complementary and Alternative Medicine. 18(1): 125 \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1186/s12906-017-1775\u0026ndash;3\u003c/span\u003e\u003cspan address=\"10.1186/s12906-017-1775\u0026ndash;3\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2018).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKim, J. Y., Choi, H. Y., Kim, H. M., Choi, J. H, \u0026amp; Jang, D. S. A Novel Cytotoxic Steroidal Saponin from the Roots of \u003cem\u003eAsparagus cochinchinensis\u003c/em\u003e. Plants. 10(10), 2067. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3390/plants10102067\u003c/span\u003e\u003cspan address=\"10.3390/plants10102067\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2021).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLiang, H. L. A. \u003cem\u003eet al\u003c/em\u003e. Effects of Miao Medicine \u003cem\u003eAsparagus\u003c/em\u003e on colorectal cancer cell proliferation and migration. Chin. J. Gerontol. 41(23), 5249\u0026ndash;5252. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://dx.doi.org/10.3969/j.issn.1005\u0026ndash;9202.2021.23.022(2021\u003c/span\u003e\u003cspan address=\"10.3969/j.issn.1005\u0026ndash;9202.2021.23.022(2021\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLiu, M. M., Zhou, Y. Z., Yuan, L. M., Shui, J. R., Xiong, L., \u0026amp; Bian, M. H. Brewing and quality analysis of \u003cem\u003eAsparagus\u003c/em\u003e new rice wine. Food Ferment. Ind. 45(1), 137\u0026ndash;144. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://dx.doi.org/10.13995/j.cnki.11\u0026ndash;1802/ts.018083\u003c/span\u003e\u003cspan address=\"10.13995/j.cnki.11\u0026ndash;1802/ts.018083\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2019).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZeng, G. P., Liu, H. C., \u0026amp; Zhang, X. Suitable Environment of \u003cem\u003eAsparagus cochinchinensis\u003c/em\u003e in Guizhou. Guizhou Agri Sci. 38, 48\u0026ndash;50. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.cabdirect.org/cabdirect/abstract/20103170869\u003c/span\u003e\u003cspan address=\"https://www.cabdirect.org/cabdirect/abstract/20103170869\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2010).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDeng, L. F., Yuan, H. R., Xie, J., Ge, L. F, \u0026amp; Chen, Y. Herbaceous plants are better than woody plants for carbon sequestration. Resources. Conserv. Recycl. 184, 106431. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.resconrec.2022.106431\u003c/span\u003e\u003cspan address=\"10.1016/j.resconrec.2022.106431\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2022).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGilardelli, F. \u003cem\u003eet al\u003c/em\u003e. Root Characteristics of Herbaceous Species for Topsoil Stabilization in Restoration Projects. Land Degrad. Dev. 28(7), 2074\u0026ndash;2085. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/ldr.2731\u003c/span\u003e\u003cspan address=\"10.1002/ldr.2731\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2017).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eYang, K. T., Han, X. M., \u0026amp; Liiang, Y. F. Effects of Planting Three Kinds of Medicinal Materials under the Forest on Soil Physicochemical Properties and Nutrients, J.Anhui Agric.Sci. 51(9): 83\u0026ndash;86. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://dx.doi.org/10.3969/j.issn.0517\u0026ndash;6611.2023.09.020\u003c/span\u003e\u003cspan address=\"10.3969/j.issn.0517\u0026ndash;6611.2023.09.020\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2023).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLiu, C. \u003cem\u003eet al\u003c/em\u003e. Plant adapt ability in karst regions. J. Plant Res. 134, 889\u0026ndash;906. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://dx.doi.org/10.1007/s10265-021-01330-3\u003c/span\u003e\u003cspan address=\"10.1007/s10265-021-01330-3\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2021).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSong, H. Y., Zhang, J., Zhao, Y. J., Teng, J., \u0026amp; Liu, J. C. Effects of rocky desertification on growth and biomass accumulation and distribution of terminal twigs in \u003cem\u003eViburnum chinshanense Graeb\u003c/em\u003en. Plant Sci. J. 36(1), 103\u0026ndash;111. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://dx.doi.org/10.11913/PSJ.2095\u0026ndash;0837.2018.10103\u003c/span\u003e\u003cspan address=\"10.11913/PSJ.2095\u0026ndash;0837.2018.10103\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2018).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZhang, X., Liu, H. C., \u0026amp; Zeng, G. P. Research on the Effect of Different Water Supply on Resistance Physiology and Appearance Charactersin \u003cem\u003eAsparagus Cochinchinensis\u003c/em\u003e (Lour.) Merr. Modern Chin. Med. 12, 19\u0026ndash;22. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://dx.doi.org/10.13313/j.issn.1673\u0026ndash;4890.2010.02.002\u003c/span\u003e\u003cspan address=\"10.13313/j.issn.1673\u0026ndash;4890.2010.02.002\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2010).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHuang, R. Y. \u003cem\u003eet al\u003c/em\u003e. Responses of Physiology, Photosynthetic Characteristics and Leaf surface Micromorphology of \u003cem\u003eBerberis poiretii\u003c/em\u003e to Drought Stress. Acta Agrestia Sin. 31, 1386\u0026ndash;1392. doi: \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.11733/j.issn.1007\u0026ndash;0435.2023.05.013\u003c/span\u003e\u003cspan address=\"10.11733/j.issn.1007\u0026ndash;0435.2023.05.013\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2023).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLiu, C. C., Liu, Y. G., Guo, K., Fan, Y., Li, G. Q., \u0026amp; Zheng, Y. R. Effect of drought on pigments, osmotic adjustment and antioxidant enzymes in six woody plant species in karst habitats of southwestern China. Environ. Exp. Bot. 71(2), 174\u0026ndash;183. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.envexpbot.2010.11.012\u003c/span\u003e\u003cspan address=\"10.1016/j.envexpbot.2010.11.012\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2011).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePan, R. Z., Wang, X. Q., \u0026amp; Li, N. H. \u003cem\u003ePlant Physiology\u003c/em\u003e, seventh ed. 81 (Higher Education Press, 2012).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHou, R. R. \u003cem\u003eet al\u003c/em\u003e. Polysaccharide from Echinacea purpurea reduce the oxidant stress in vitro and in vivo. Int. J. Biol. Macromol. 149, 41\u0026ndash;50. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.ijbiomac.2020.01.129\u003c/span\u003e\u003cspan address=\"10.1016/j.ijbiomac.2020.01.129\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2020).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHuang, H. L., Chen, F., Long, R., \u0026amp; Huang, J. L. The antioxidant activities in vivo of bitter gourd polysaccharide. J. Biol. Macromol. 145, 141\u0026ndash;144. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.ijbiomac.2019.12.165\u003c/span\u003e\u003cspan address=\"10.1016/j.ijbiomac.2019.12.165\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2020).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLin, Y. Y. \u003cem\u003eet al\u003c/em\u003e. Enzyme and microwave coassisted extraction, structural characterization and antioxidant activity of polysaccharides from Purple-heart Radish. Food Chemistry. 372, 131274. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://dx.doi.org/10.1016/j.foodchem.2021.131274\u003c/span\u003e\u003cspan address=\"10.1016/j.foodchem.2021.131274\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2022).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWan, L. Y., Fu, J. E., Pan, L. M., Ke, F., \u0026amp; Wei, S. G. Analysis and evaluation of agronomic traits of Guangxi \u003cem\u003eAsparagus\u003c/em\u003e germplasm resources. Jiangsu Agri Sci. 48, 124\u0026ndash;128. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://dx.doi.org/10.15889/j.issn.1002\u0026ndash;1302.2020.24.023\u003c/span\u003e\u003cspan address=\"10.15889/j.issn.1002\u0026ndash;1302.2020.24.023\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2020).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePeng, X., Huang, Y., Che, J. X., \u0026amp; Lu, L. Changing rule of soil physicochemical features under different grade of desertification A case study at Puding station of Guizhou karst center. China Karst. 28(4): 402\u0026ndash;405. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://dx.doi.org/doi:10.3969/j.issn.1001\u0026ndash;4810.2009.04.012\u003c/span\u003e\u003cspan address=\"doi:10.3969/j.issn.1001\u0026ndash;4810.2009.04.012\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2009).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eViviana, L. P., Engil, P. P., Rapha\u0026euml;l, W., Marcel, H., \u0026amp; Johan, S. Improvement of soil structure through organic crop management, conservation tillage and grass-clover ley. Soil Tillage Res. 180, 1\u0026ndash;9. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://dx.doi.org/10.1016/j.still.2018.02.007\u003c/span\u003e\u003cspan address=\"10.1016/j.still.2018.02.007\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2018).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eManlay, R. J., Feller, C., \u0026amp; Swift, M. J. Historical evolution of soil organic matter concepts and their relationships with the fertility and sustainability of cropping systems. Agric, Ecosyst. Environ. 119, 217\u0026ndash;233. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.agee.2006.07.011\u003c/span\u003e\u003cspan address=\"10.1016/j.agee.2006.07.011\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2007).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWu, Y. N., Yu. L. F., Zhang, L. M., Liu, N., \u0026amp; Yan, L. B. Characteristics and influencing factors of soil carbon pool during vegetation restoration in Karst Plateau. Ecol. Environ. Sci. 29, 1935\u0026ndash;1942. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.jeesci.com/EN/Y2020/\u003c/span\u003e\u003cspan address=\"https://www.jeesci.com/EN/Y2020/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003eV29/I10/1935 (2020).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZhang, W., Zhao, J., Pan, F., Li, D. J., \u0026amp; Chen, H. S. Changes in nitrogen and phosphorus limitation during secondary succession in a karst region in southwest China. Plant Soil. 391, 77\u0026ndash;91. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://dx.doi.org/10.1007/s11104-015-2406\u0026ndash;8\u003c/span\u003e\u003cspan address=\"10.1007/s11104-015-2406\u0026ndash;8\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2015).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePang, L., Wu, X., Zou, Q. L., \u0026amp; Li, J.X. Distribution of N and P of Common Ferns and their Relationship with Soil Supply in Qianzhong Karst Area. J Northeast For Univ. 49, 53\u0026ndash;57. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://dx.doi.org/10.13759/j.cnki. dlxb.2021.06.011\u003c/span\u003e\u003cspan address=\"10.13759/j.cnki. dlxb.2021.06.011\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2021).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMing, X., \u0026amp; Chen, W. N. Stomata Distribution in Leaves of Different Plants. Anhui Agri Sci Bull. 27(20), 22\u0026ndash;25. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://dx.doi.org/10.16377/j.cnki.issn1007\u0026ndash;7731.2021.20.009\u003c/span\u003e\u003cspan address=\"10.16377/j.cnki.issn1007\u0026ndash;7731.2021.20.009\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2021).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLi, H. S. \u003cem\u003ePrinciples of plant physiological and biochemical experimental techniques\u003c/em\u003e. 134\u0026ndash;137. (Higher Education Press, 2000).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLi, M., Fei, Y., Li, L. X., \u0026amp; Fang, D. S. Method Research to Assay Total Saponin Contentin \u003cem\u003eAsparagus Cochinchinese\u003c/em\u003e Root. J. Chengdu Univ TCM. 4, 46\u0026ndash;48. (2001).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWu, F., Liu, M. D., Li, Y. X., \u0026amp; Jin, C.S. Study on Extraction Technology of Total Amino Acids and Polysaccharides from \u003cem\u003eAsparagus\u003c/em\u003e. J Chizhou Univ. 31(3), 60\u0026ndash;63. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://dx.doi.org/10.13420/j.cnki.jczu\u003c/span\u003e\u003cspan address=\"10.13420/j.cnki.jczu\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e. 2017.03.0021 (2017).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBao D. S. \u003cem\u003eSoil Agrochemical Analysis\u003c/em\u003e. third ed. 14\u0026ndash;118. (Beijing Agricultural Press, 2013).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHu, Y., Wang, Y., \u0026amp; Li, G. Soil Nutrient Characteristics and Fertility Grade Evaluation of Newly-increased Farmland in Weibei Dryland: A Case Study of Heyang County. Chin. Agri. Sci Bull. 38, 94\u0026ndash;100. doi:\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.11924/j.issn.1000\u0026ndash;6850.casb2021\u0026ndash;0886\u003c/span\u003e\u003cspan address=\"10.11924/j.issn.1000\u0026ndash;6850.casb2021\u0026ndash;0886\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2022).\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Asparagus munitus, karst rocky desertification, agronomic traits, soil physical properties, soil nutrients","lastPublishedDoi":"10.21203/rs.3.rs-4318709/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-4318709/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eIn this study, \u003cem\u003eAsparagus munitus\u003c/em\u003e was planted in karst rocky desertification (KRD) areas and arable land (AL) to explore the changes in plant adaptability to KRD environments and effects on KRD soils. Results showed \u003cem\u003eA. munitus\u003c/em\u003e exhibited shrinkage in KRD habitats, evidenced by increased number and length of stems, branch length, and other agronomic traits; significantly elevated antioxidant enzyme and osmoregulator content in leafy shoots; and increased polysaccharide content in tuberous roots. Correlation and principal component analyses of plant agronomic traits revealed strong correlations among indicators except basal stems, with leaf shoot length and root diameter as the first and second principal components, respectively. Soil physical properties and nutrient contents showed KRD soil pH, water content, porosity, and nutrient content markedly changed after \u003cem\u003eA. munitus\u003c/em\u003e planting, overall positively developing. This study reveals partial adaptive strategies of \u003cem\u003eA. munitus\u003c/em\u003e to KRD habitats and ameliorative effects of planting \u003cem\u003eA. munitus\u003c/em\u003e on KRD soils, providing theoretical bases for \u003cem\u003eA. munitus\u003c/em\u003e cultivation and KRD soil improvement.\u003c/p\u003e","manuscriptTitle":"Adaptation of Asparagus munitus to karst adversity and its effects on soil","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-05-02 04:09:34","doi":"10.21203/rs.3.rs-4318709/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":"cba8f331-41f5-4939-884b-e512bcbce0ce","owner":[],"postedDate":"May 2nd, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[{"id":31322264,"name":"Biological sciences/Physiology"},{"id":31322265,"name":"Biological sciences/Plant sciences"},{"id":31322266,"name":"Earth and environmental sciences/Environmental sciences"}],"tags":[],"updatedAt":"2024-10-03T03:38:14+00:00","versionOfRecord":[],"versionCreatedAt":"2024-05-02 04:09:34","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-4318709","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-4318709","identity":"rs-4318709","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}