Effects of cadmium pollution on soil feedback between invasive plant Phytolacca Americana and native plant Phytolacca Acinosa

Effects of cadmium pollution on soil feedback between invasive plant Phytolacca Americana and native plant Phytolacca Acinosa | Authorea try { document.documentElement.classList.add('js'); } catch (e) { } var _gaq = _gaq || []; _gaq.push(['_setAccount', 'G-8VDV14Y67G']); _gaq.push(['_trackPageview']); (function() { var ga = document.createElement('script'); ga.type = 'text/javascript'; ga.async = true; ga.src = ('https:' == document.location.protocol ? 'https://ssl' : 'http://www') + '.google-analytics.com/ga.js'; var s = document.getElementsByTagName('script')[0]; s.parentNode.insertBefore(ga, s); })(); Skip to main content Preprints Collections Wiley Open Research IET Open Research Ecological Society of Japan All Collections About About Authorea FAQs Contact Us Quick Search anywhere Search for preprint articles, keywords, etc. Search Search ADVANCED SEARCH SCROLL This is a preprint and has not been peer reviewed. Data may be preliminary. 21 February 2025 V1 Latest version Share on Effects of cadmium pollution on soil feedback between invasive plant Phytolacca Americana and native plant Phytolacca Acinosa Authors : Yue Zhu 0009-0009-3207-2870 , Jun Chen , Yu Ming , Jingru Zhang , Shaoyu Zhang , Yunshan Liu , Bo Li , Jihua Wu 0000-0001-8623-8519 , Evan Siemann , and Yi Wang 0000-0003-1851-8994 [email protected] Authors Info & Affiliations https://doi.org/10.22541/au.174013593.35371378/v1 Published Journal Of Plant Ecology Version of record Peer review timeline 227 views 147 downloads Contents Abstract 1. INTRODUCTION 2. Material and Methods Supplementary Material References Information & Authors Metrics & Citations View Options References Figures Tables Media Share Abstract Background and Aims: Soil abiotic and biotic properties shaped by plants impact future plant invasions (“plant-soil feedbacks”). Human activities have increased soil heavy metal concentrations, often where invasion pressure is increasing, so it is imperative to investigate plant-soil feedback mechanisms associated with invasive species in environments contaminated by heavy metals. Methods: We experimentally created soils with different Cd concentrations and conditioned them with invasive Phytolacca americana or native Phytolacca acinosa (or without plants) in a greenhouse experiment. We measured soil chemical, microbial (by PFLAs), and physical properties. We grew individual plants of P. Americana or P. Acinosa in these soils in a feedback phase. Results: Soil Cd increased invasive mass but decreased native mass. Fungi and bacteria were abundant in invasive conditioned soils, especially without Cd. Phenol, flavonoid and tannin concentrations and soil pH increased with Cd when plants were present. Phenol and tannin concentrations were higher with invasive plants but flavonoids were lower. In the feedback phase, invasive mass was sensitive to soil microbes. Native plants were smaller in invasive (away) soils, especially with higher soil Cd concentrations, reflecting their sensitivity to soil chemicals and soil physical characteristics. Conclusion: These results indicate that P. Americana invasions are enhanced both through direct effects of soil Cd on P. Acinosa as well as through Cd magnifying the negative effects of soil chemical and physical characteristics on the native. This study offers valuable insights into the influence of PSF on plant invasion processes in the context of soil heavy metal pollution. Effects of cadmium pollution on soil feedback between invasive plant Phytolacca Americana and native plant Phytolacca Acinosa [Redacted for peer-review] Abstract Background and Aims: Soil abiotic and biotic properties shaped by plants impact future plant invasions (“plant-soil feedbacks”). Human activities have increased soil heavy metal concentrations, often where invasion pressure is increasing, so it is imperative to investigate plant-soil feedback mechanisms associated with invasive species in environments contaminated by heavy metals. Methods: We experimentally created soils with different Cd concentrations and conditioned them with invasive Phytolacca americana or native Phytolacca acinosa (or without plants) in a greenhouse experiment . We measured soil chemical, microbial (by PFLAs), and physical properties. We grew individual plants of P. Americana or P. Acinosa in these soils in a feedback phase. Results: Soil Cd increased invasive mass but decreased native mass. Fungi and bacteria were abundant in invasive conditioned soils, especially without Cd. Phenol, flavonoid and tannin concentrations and soil pH increased with Cd when plants were present. Phenol and tannin concentrations were higher with invasive plants but flavonoids were lower. In the feedback phase, invasive mass was sensitive to soil microbes. Native plants were smaller in invasive (away) soils, especially with higher soil Cd concentrations, reflecting their sensitivity to soil chemicals and soil physical characteristics. Conclusion: These results indicate that P. Americana invasions are enhanced both through direct effects of soil Cd on P. Acinosa as well as through Cd magnifying the negative effects of soil chemical and physical characteristics on the native. This study offers valuable insights into the influence of PSF on plant invasion processes in the context of soil heavy metal pollution. Keywords: Phytolacca americana , heavy metal Cd, plant invasion, plant-soil feedback, soil biota, soil secondary metabolites 1. INTRODUCTION The global economic losses and management costs associated with biological invasions exceed $1.288 trillion per year (Diagne et al ., 2021). Plant invasions significantly change plant communities globally and are a major contributor to biodiversity loss (Broadbent et al. , 2018; Weidlich et al ., 2020). A critical area of investigation within invasion ecology is the interaction between invasive species and native plants, especially through plant-soil feedback mechanisms. Plant-soil feedback is essential for elucidating community succession and plant invasion dynamics because it links current and future plant communities (Bever et al. , 2003; Bennett et al. , 2017; Manya et al. , 2020). Plant-soil feedback (PSF) pertains to the alteration of both biological and abiotic properties of soil by plants, which subsequently influences the performance of the same or different plant species in that soil (van der Putten et al ., 2013; Crawford et al ., 2019). The mechanisms underlying PSF encompass various factors, including soil pH, microorganisms, and secondary chemicals. For instance, soil pathogens have been shown to be a key factor in negative PSF because they have limited host ranges (Kempel et al ., 2018). In contrast, the enrichment of beneficial microorganisms in the rhizosphere soil may create positive PSFs (Meng et al ., 2022). Furthermore, secondary metabolites present in the soil can also influence the growth of plants (Ning et al. , 2016). Depending on the specificity of chemical effects, secondary metabolites may limit conspecifics when they are autotoxic, but they may favor conspecifics when they primarily have allelopathic effects on other plants (Zhou et al ., 2018). When the net effect of changes in soil properties favors invasive plants over natives, it may increase invasions (van der Putten et al ., 2013). Current studies have found that negative plant-soil feedback plays an important role in maintaining plant species diversity, while positive plant-soil feedback may lead to community homogenization, especially in the context of invasions (Bever, 2003; Bever et al ., 2010). Increasing heavy metal pollution has emerged as a significant abiotic stressor impacting plant growth and development (Parmar et al ., 2013). Consequently, researchers have directed their attention toward understanding how abiotic factors, particularly heavy metal pollution, influence plant invasion and expansion (Dai et al ., 2020; Li et al ., 2021). Cadmium (Cd), a heavy metal known for its high toxicity (Ma et al. , 2021), has been continuously released and accumulated in the soil due to various natural and anthropogenic activities (Edreva et al. , 2008), resulting in severe pollution issues in China (Huang et al. , 2019). In the context of global environmental change, alterations in various environmental factors, such as nitrogen deposition, drought events, and temperature fluctuations, impose selection pressures on ecological communities and influence plant-soil feedback mechanisms within these communities (van der Putten et al ., 2016). Total phenolic compounds are frequently used to indicate secondary metabolic allelochemicals in plants (Lou et al. , 2016). Additionally, flavonoids play a significant role in various processes, including antioxidant and antibacterial activities (Daglia, 2012). Furthermore, tannins are commonly employed for removing heavy metal ions (Xu et al. , 2017), and their presence has been shown to mitigate the toxicity of these metal ions (Qiao et al. , 2021). In general, heavy metals might change the production of secondary compounds released into the soil or may have different effects on plants depending on heavy metal stress. Furthermore, soil heavy metal pollution serves as a critical selective stressor, which has the potential to modify the dynamics of plant-soil feedback (Zhang et al ., 2007). However, there is a lack of research examining plant-soil feedback in soils contaminated by heavy metals, and the interactions among plants, soil, and heavy metal pollution remain inadequately understood. Phytolacca americana (Phytolaccaceae) is a perennial herb native to North America that was introduced to China approximately 90 years ago (Xu et al. , 2006). It is toxic, posing direct risks to humans and livestock (Zhang et al. , 2014). It has the ability to hyperaccumulate the heavy metal cadmium (Cd) (Zhao et al. , 2022) and natives enhance its growth in open areas via changes in soils (Chen et al. , 2021). Consequently, P. Americana has established a competitive advantage over native flora (Xiao et al. , 2019), leading to its continued spread that causes considerable ecological damage in various regions of China (Huang et al. , 2016). Phytolacca acinosa , a native congener of the invasive P. Americana in China, shares similar distribution and resource requirements. This study evaluated the following hypotheses: (1) Will Cd increase invasions by increasing the sensitivity of a native plant to soil conditions created by an invasive plant and/or by decreasing the sensitivity of an invasive plant to soil conditions created by a native plant? (2) Will Cd increase invasions by changing the magnitude of effects of the native and/or invasive on the soil? (3) Are the Cd-dependent effects of the invasive on the native (and vice versa) mediated through changes in soil chemistry, soil physical properties, and/or soil microbes? 2. Material and Methods 2.1 Plant materials In May 2021, we collected P. Americana seeds from an unpolluted area near Yunnan University in Kunming, Yunnan, China (24.89°N, 102.82°E) and P. Acinosa seeds from an unpolluted area in Qujing, Yunnan (25.27°N, 104.19° E). We surface sterilized the P. Americana and P. Acinosa seeds with a 7.2% sodium hypochlorite solution for 5 minutes. Subsequently, we immersed the P. Americana and P. Acinosa seeds in 98% concentrated sulfuric acid for 6 minutes and 9 minutes, respectively, to facilitate the breaking of dormancy then rinsed with water. Then, we transferred the seeds to an incubator set at 25°C, with a photoperiod of 16 hours of light per day and a relative humidity of 65% to promote germination. Approximately one-week post-germination, we planted the seedlings to a container filled with commercial humus soil. Once the plants developed a pair of cotyledons, we selected healthy seedlings exhibiting consistent growth for use in the conditioning and soil feedback phases of the experiment. 2.2 Soil collection The background soil utilized in this study was commercial humus soil. We collected the soil inoculum in October 2022 from farmland in Kunming, where P. Americana and P. Acinosa were not present (24.85°N, 102.94°E). We removed the surface litter and then the surface soil (0-10 cm) was collected, dried, screened, and stored at room temperature for the experiments. 2.3 Experimental design We conducted a greenhouse experiment to investigate the PSFs of the invasive plant P. Americana and its native congener P. Acinosa in the context of cadmium (Cd) pollution (Fig. 1). First, we created soils with four levels of Cd contamination. Second, we grew the two Phytolacca species in these four soils or left soils without plants in the soil conditioning phase (144 pots: 4 Cd concentrations × 3 conditioned plant species × 12 replicates). Third, in a factorial design, we tested the growth of the two Phytolacca species in the conditioned soil types that were sterilized or treated with activated charcoal. The soil feedback phase consisted of 1152 pots (144 conditioning phase soils × four soil treatments × 2 responding species). 2.4 Soil preparation phase In November 2022, we disinfected 144 twenty-six-liter pots and trays using a 7.2% sodium hypochlorite solution and then rinsed them. We filled each pot with 8.5 kg of a mixture of commercial soil (97% by weight) and field-collected soil inoculum (3% by weight). Based on prior research and the current state of soil heavy metal pollution (Garg et al ., 2012; Duan et al. , 2016; Dou et al. , 2022), we created four soil concentrations of cadmium (Cd) (0, 2, 8, 32 mg Cd/kg). We applied 20 ml of solutions with different concentrations of CdCl 2 ·2.5H 2 O or distilled water (36 pots for each concentration), thoroughly mixed the soils and allowed them to equilibrate for two weeks in a greenhouse at Yunnan University (24.83° N, 102.84° E) under controlled environmental conditions (40/14°C, 12 hours of light per day, and 45–70% relative humidity) following Wang et al. , 2021. 2.5 Soil conditioning phase We assigned each pot to a conditioning treatment ( P. Americana, P. Acinosa, or no plants) . We left pots unplanted, or we transplanted four seedlings of P. Americana or P. Acinosa into each pot and replaced any seedlings that died in the first two weeks. Then we thinned pots to three seedlings. We watered pots every 2 to 3 days to keep them moist and randomly exchanged the positions of the pots twice a month. We left plants growing from December 2022 to May 2023 (~five months). We measured plant height, clipped them at ground level, dried them (60°C for 48 hours) and weighed the aboveground biomass. We removed root systems from the pots and collected the rhizosphere soil adhering to the roots by gently shaking them (Wang et al ., 2020). We preserved some of the soil at -20°C for future analysis of soil microbiological properties. We stored another portion at 4°C for subsequent assessment of physical and chemical properties. The rest of the soil was air-dried and then stored in plastic bags for the feedback phase 20 days later. Without plant transpiration, the potential soil moisture loss between the two phases was deemed negligible (Koyama et al. , 2022). Then we washed, dried, and weighed the roots. 2.6 Soil feedback phase We filled 288 three-L pots with treated soils. We germinated seeds of P. Americana and P. Acinosa following the same methods as for the conditioning phase. We transplanted two seedlings of P. Americana or P. Acinosa into each pot and replaced any seedlings that died in the first two weeks. Then we thinned pots to a single seedling. We watered pots every 2 to 3 days to keep them moist, checked survival and randomly exchanged the positions of the pots twice a month. We left plants growing from June 2023 to October 2023 (~five months). We measured plant height then clipped them at ground level then dried (60°C for 48 hours) and weighed the aboveground biomass. Then we washed, dried, and weighed the roots. 2.7 Determination of Soil Properties We assessed soil bulk density for four replicates of each conditioning treatment combination using the ring knife method and soil water content through the drying method. We sieved soil samples through a 10-mesh sampling screen and then measured pH through a suspension of soil in deionized water (Zhang et al ., 2020). We determined soil electrical conductivity (EC) using the same soil-deionized water suspension method (Qiao et al. , 2021). 2.8 Determination of soil secondary metabolites We extracted and quantified total phenols in soil for four replicates of each conditioning treatment combination (same ones as for pH) following Sahayarayan et al. (2020) and Nicholas et al . (2021). We extracted 10 g of soil using 100 mL of 95% ethanol and concentrated it to approximately 2 mL using a rotary evaporator. Then, we combined 0.5 mL of the sample with 2 mL of Folin-Ciocalteu reagent and 4 mL of a 7.5% sodium carbonate solution and incubated the mixtures for 30 minutes at 23°C. We measured the absorbance at 765 nm with a gallic acid standard. We quantified total flavonoids in the soil for four replicates of each conditioning treatment combination using a modified colorimetric method (Phares et al ., 2020; Wang et al ., 2020). We extracted 10 grams of soil with methanol and measured absorbance at 510 nm with catechin as the standard. We quantified the concentration of tannins for four replicates of each conditioning treatment combination in the soil following Qiao et al. (2021). We extracted a 2 g sample of sieved soil (2 mm mesh) with methanol and then centrifuged the mixture at 5000 rpm for 10 minutes to obtain the supernatant. Subsequently, we added 6 mL of a 4% vanillin methanol solution and 3 mL of concentrated hydrochloric acid to a 25 mL volumetric flask shielded from light. We allowed it to react at 20°C for 15 hours. We measured absorbance at 510 nm with catechin as the standard. 2.9 Determination of soil microbiological properties We estimated microbial biomass on three of the four replicates with plants (i.e., no plant-free pots) used for measuring soil secondary metabolites using soil phospholipid fatty acids (PLFA) following Spitzer et al. (2021). We extracted PLFAs from 4 grams of freeze-dried soil, utilizing C19:0 as the internal standard. We quantified PLFAs using an Agilent 7890B gas chromatograph equipped with a flame ionization detector (GC-FID). We used the MIDI Sherlock software system to identify and characterize the PLFAs. We used the following fatty acids as indicators for bacterial presence: i14:0, 14:0, i-15:0, a-15:0, 15:0, i-16:0, 16:1ω9, 16:1ω7c, 16:1ω7t, i-17:0, a-17:0, 17:1ω8, cy-17:0, 17:0, 18:1ω7, and cy-19:0 (Spitzer et al ., 2021). We used 18:1ω9c and 18:2ω6c as indicators for fungal presence (Zhang et al. , 2016). For the identification of gram-positive bacteria, we used i-15:0, a-15:0, i-16:0, i-17:0, and a-17:0, whereas we used cy-17:0, 18:1ω7, and cy-19:0 as indicators for gram-negative bacteria (Wardle et al ., 2013). We calculated the absolute abundance of PLFAs in micrograms per gram of soil. We used the PLFA data to calculate the ratios of fungi to bacteria and gram-positive to gram-negative bacteria. Analyses were conducted at Hangzhou Yanqu Information Technology Co., Ltd. 2.10 Statistical analysis We used ANOVAs to determine the dependence of plant height and mass (total mass, aboveground mass, belowground mass, root: shoot), soil physical properties (pH, electrical conductivity, bulk density, water content), soil chemical properties (phenols, flavonoids, tannins), and soil microbes (fungi: bacteria, fungi, bacteria, gram-positive bacteria, gram-negative bacteria, gram-positive: gram-negative) on Cd concentrations (0, 2, 8, 32 mg/kg), plant treatment ( P. Americana , P. Acinosa , no plants) and their interaction in ANOVAs. We used adjusted means post-hoc tests to examine the differences among means for factors with more than two levels. We used Spearman correlations to test the relationships of invasive plant mass and native plant mass in the feedback phase on Cd treatment (as a continuous variable), conditioning plant mass, phenols, flavonoids, tannins, fungi: bacteria, fungi, bacteria, gram-positive bacteria, gram-negative bacteria, gram-positive: gram-negative, soil pH, soil electrical conductivity, soil bulk density, and soil water content. We used mixed model ANOVA to test the dependence of mass in the feedback phase on the fixed factors Cd treatment (0, 2, 8, 32 mg/kg), conditioning origin (invasive or native), feedback origin (invasive or native) and the interactions plus the random factor soil (Cdtrt×CondO) in an ANOVA. We calculated each replicate’s PSF index as ln(home/away) for each conditioning phase. We calculated the 95% confidence intervals. We considered intervals that did not overlap zero to be significantly different than zero and intervals that did not overlap to be significantly different. 3. RESULTS 3.1 Conditioning phase In the conditioning phase, plant height (Fig. S1) plus total (Fig. 1A), aboveground (Fig S1B) and belowground mass (Fig. S1D) all depended on the interaction of Cd treatment and species origin (Table 1). At 0 mg/kg Cd, their masses were comparable. Phytolacca americana height and mass increased from 2 to 8 mg/kg Cd, then decreased to be similar to the 0 mg/kg treatment at 32 mg/kg. In contrast, P. Acinosa height and mass decreased from 8 to 32 mg/kg Cd so that its mass was lower than P. Americana . Soil pH was higher when soil was conditioned by P. Americana (5.01±0.03) than P. Acinosa (4.89±0.07) but lower than plant-free pots (5.21±0.03; Table 1, Fig. 2D). Soil pH, EC, and bulk density decreased with soil Cd concentration (Table 1, Fig. 2DEF). SWC did not depend on any conditioning treatments (Table 1, Fig. S2D). Phenols and tannins in soil depended on Cd treatment × conditioning plant species with low concentrations in soils conditioned without plants for every Cd concentration, higher concentrations in P. Americana conditioned soils than P. Acinosa conditioned soils, and increasing concentrations with Cd concentrations in the soils conditioned by either plant species (Table 1, Fig. 2AC). Flavonoids did not vary with Cd concentration, but they were low in soils conditioned without plants (1.60±0.04 mg/kg), intermediate in P. Americana conditioned soils (2.03±0.03 mg/kg), and high in P. Acinosa conditioned soils (2.13±0.13 mg/kg; Table 1, Fig 2B). Fungi: bacteria is higher in P. Americana conditioned soils than in P. Acinosa conditioned soils, and it decreased with Cd concentration (Table 1, Fig. S2B). Fungi and bacteria (all gram positive, gram negative) depended on Cd treatment × conditioning plant species. Fungi were more abundant in 0 Cd, P. Americana soils, intermediate in other P. Americana conditioned soils and 0 Cd P. Acinosa soils, and less abundant in P. Acinosa soils with Cd (Fig. 1B). Bacteria were higher in low Cd P. Americana conditioned soils, low in 8 mg/kg Cd soils, and intermediate in other Cd and plant species combinations (Fig. 2CD, S2A). The gram-positive to gram-negative bacteria ratio did not depend on treatments (Table 1, Fig. S2C). 3.2 Feedback phase In the feedback phase, total mass, aboveground mass and belowground mass of both P. Americana and P. Acinosa decreased with Cd concentrations (Table 2, Fig 3A, Fig. S3AB). These variables also all depended on the interaction of conditioning and feedback origins (Table 2). This reflected a similar mass of P. Americana in soils conditioned by either species or without plants but a lower mass of P. Acinosa in soils conditioned by P. Americana (Fig. 3A, Fig. S3AB). Because the magnitude of changes in aboveground and belowground mass differed, root: shoot depended on the three-way interaction of Cd trt × conditioning plant × feedback origin (Table 2). Both species had the lowest R:S in soils with 0 Cd conditioned without plants, higher R:S in 0 Cd with either plant in the conditioning stage, and increasing R:S with Cd concentration, but the increase with Cd was especially large for P. Americana in P. Americana conditioned soils (“home soils, Fig. 3B). The PSF index for P. Americana did not differ from 0 except for in 8 mg/kg Cd soils, for which it was slightly positive (i.e., better performance in home soil; Fig. 4). The PSF for P. Acinosa was always positive, and it increased with Cd concentration (Fig. 4). The mass of both species was negatively correlated with the concentrations of phenols, flavonoids, and tannins in the soil. However, the correlations were more substantial for P. Acinosa compared to P. Americana (Fig. 5). The mass of P. Americana was negatively correlated with bacteria and gram-negative bacteria. However, the mass of P. Acinosa was not correlated with either. The mass of both species was positively correlated with soil pH, but the correlation was stronger for P. Acinosa . The mass of P. Acinosa was positively correlated with soil EC and SWC, but the mass of P. Americana was not. Other variables were not correlated with the mass of either species (Fig. S4). 4. Discussion This study examined the plant-soil feedback (PSF) mechanisms of P. Americana and P. Acinosa under varying concentrations of cadmium (Cd) to elucidate the competitive advantage of invasive species in polluted environments. The results demonstrate that increasing Cd concentrations enhanced the positive PSF of the invasive P. Americana while amplifying the negative PSF of the native P. Acinosa. These findings reinforce the notion that PSF dynamics can play a critical role in facilitating the success of invasive species under environmental stressors, such as heavy metal contamination.The rhizosphere soil properties of P. Americana and P. Acinosa were significantly influenced by Cd concentrations, with important implications for their feedback dynamics. Consistent with previous studies (Li et al., 2023), the pH of rhizosphere soils decreased as Cd concentrations increased. However, P. Americana maintained higher pH levels compared to P. Acinosa, suggesting a potential mechanism by which the invasive species mitigates the toxicity of available Cd. This observation aligns with evidence that elevated pH can reduce Cd bioavailability (Janousková et al., 2010), thereby diminishing its deleterious effects on plant growth and promoting the competitive dominance of invasive species.Secondary metabolites also emerged as critical factors shaping PSF dynamics. Higher concentrations of phenols and tannins were observed in the rhizosphere soils of P. Americana at increased Cd levels, consistent with the hypothesis that invasive species modulate soil chemistry to mitigate stress and outcompete native plants (Qiao et al., 2021). Phenolics and tannins are known to interact with heavy metals, potentially reducing their toxicity (Xu et al., 2017). While flavonoids exhibited distinct patterns, the overall accumulation of secondary metabolites in P. Americana-conditioned soils underscores the role of chemical interactions in driving PSF outcomes.Microbial community shifts further contributed to the observed feedback effects. The abundance of fungi and bacteria declined with increasing Cd concentrations in the soils of both species, a trend consistent with prior research on the impacts of heavy metal stress on microbial communities (Marchante et al., 2008; Stefanowicz et al., 2016). However, the fungi-to-bacteria ratio remained consistently higher in P. Americana-conditioned soils, suggesting that fungi, which are closely associated with plant roots, may enhance the resilience of invasive species in stressful environments (Drigo et al., 2010). Although soil biota exerted an inhibitory effect on the growth of P. Americana, this effect diminished as Cd concentrations increased, indicating a potential adaptive response of the invasive species to microbial pressures in contaminated soils.Overall, the findings suggest that the interaction between altered soil chemistry, secondary metabolites, and microbial communities creates feedback loops that favor P. Americana in Cd-contaminated environments. These results highlight the complexity of PSF mechanisms and their role in shaping plant-plant interactions in polluted ecosystems. Nonetheless, the study is limited by its lack of temporal analysis. Future research should incorporate longitudinal studies to examine how PSF dynamics evolve over time, particularly as soil contamination persists or intensifies. 5. Conclusion This study provides novel insights into the role of plant-soil feedback (PSF) mechanisms in mediating the competitive interactions between the invasive Phytolacca americana and the native Phytolacca acinosa under cadmium-contaminated conditions. The findings reveal that increasing Cd concentrations positively influenced the PSF of P. Americana while exacerbating the negative PSF of P. Acinosa. These patterns suggest that PSF dynamics are critical in facilitating the establishment and persistence of invasive species in polluted ecosystems. Key drivers of these feedback effects included differences in soil pH, secondary metabolite accumulation, and microbial community composition. The higher pH and increased accumulation of phenols and tannins in P. Americana-conditioned soils likely conferred a competitive advantage to the invasive species by reducing Cd toxicity and altering the soil environment to its benefit. The higher fungi-to-bacteria ratio in P. Americana soils further supported its adaptive advantage under heavy metal stress. Despite these advances, the study highlights the need for further research to address its limitations. In particular, the lack of temporal analyses precludes understanding how PSF dynamics may shift over time as invasive species establish or as soil contamination progresses. Longitudinal studies that integrate temporal dimensions and examine different stages of invasion are necessary to provide a more comprehensive understanding of PSF processes. In conclusion, this study underscores the importance of PSF mechanisms in shaping the ecological success of invasive species in polluted environments. By elucidating how invasive plants exploit altered soil properties, microbial interactions, and chemical processes, these findings contribute to the broader understanding of invasion ecology and offer potential strategies for managing invasive species and restoring degraded ecosystems Acknowledgment [Redacted for peer-review] Cd Plant Cd × Plant Variable df F P df F P df F P Plants Height 3,88 15.88 <.0001 1,88 64.51 <.0001 3,88 6.51 0.0005 Total mass 3,88 5.07 0.0028 1,88 22.10 <.0001 3,88 3.20 0.0274 AG mass 3,88 3.73 0.0142 1,88 36.14 <.0001 3,88 3.35 0.0227 BG mass 3,88 7.12 0.0002 1,88 10.60 0.0016 3,88 2.89 0.0399 R:S 3,88 22.07 <.0001 1,88 75.02 <.0001 3,88 1.12 0.3442 Soil physical properties pH 3,36 14.02 <.0001 2,36 21.14 <.0001 6,36 1.12 0.3691 EC 3,36 1.71 0.1819 2,36 4.71 0.0152 6,36 0.47 0.8253 Bulk Density 3,36 0.37 0.7759 2,36 6.96 0.0028 6,36 0.37 0.8922 SWC 3,36 0.48 0.6985 2,36 0.55 0.5838 6,36 0.07 0.9983 Soil chemical properties Phenols 3,36 7.90 0.0004 2,36 439.11 <.0001 6,36 2.44 0.0441 Flavonoids 3,36 1.58 0.2109 2,36 12.73 <.0001 6,36 0.78 0.5885 Tannins 3,36 12.24 <.0001 2,36 144.03 <.0001 6,36 4.66 0.0013 Soil microbes Fungi: Bacteria 3,16 4.89 0.0134 1,16 7.69 0.0136 3,16 0.50 0.6854 Fungi 3,16 14.43 <.0001 1,16 21.79 0.0003 3,16 3.45 0.0418 Bacteria 3,16 14.18 <.0001 1,16 21.42 0.0003 3,16 20.69 <.0001 Gram + 3,16 4.69 0.0155 1,16 3.31 0.0877 3,16 14.22 <.0001 Gram - 3,16 8.09 0.0017 1,16 11.93 0.0033 3,16 11.34 0.0003 + to - 3,16 1.15 0.3604 1,16 1.05 0.3198 3,16 1.75 0.1975 Table 2. Feedback phase. The dependence of mass in the feedback phase on the fixed factors Cd treatment (0, 2, 8, 32 mg/kg), conditioning origin (invasive or native), feedback origin (invasive or native), and the interactions plus the random factor soil (Cdtrt×CondO) in an ANOVA. Significant results are indicated in bold (P<0.05). Total mass AG mass BG mass R:S Effect df F P F P F P F P Cd treatment 3,36 10.33 <.0001 18.8 <.0001 5.47 0.0034 257.08 <.0001 Conditioning origin 2,36 24.01 <.0001 28.14 <.0001 20.20 <.0001 26.99 <.0001 Cdtrt×CondO 6,36 1.91 0.1059 1.82 0.1232 2.02 0.0882 4.98 0.0008 Feedback origin 1,228 150.52 <.0001 138.36 <.0001 160.26 <.0001 44.51 <.0001 Cdtrt×FBO 3,228 2.00 0.1153 1.80 0.1484 2.37 0.0713 6.43 0.0003 CondO×FBO 2,228 23.9 <.0001 19.03 <.0001 28.49 <.0001 41.88 <.0001 Cdtrt×CondO×FBO 6,228 1.56 0.1600 1.14 0.3398 2.04 0.0611 7.08 <.0001 Figure 1. The dependence of (A) plant mass and soil abundance of (B) soil fungi, (C)soil gram-positive bacteria, and (D) gram-negative bacteria on soil Cd concentrations and plant species in the conditioning phase. Means ± 1 s.e. Letters indicate means that did not differ in post hoc tests (P<0.05). Figure 2. The dependence of soil concentrations of (A) phenols, (B) flavonoids, and (C) tannins, (D) soil pH, (E) soil electrical conductivity, and (F) soil bulk density on soil Cd concentrations and plant treatment in the conditioning phase. Means ± 1 s.e. Letters indicate means that did not differ in post hoc tests (P<0.05). Figure 3. The dependence of plant (A) mass and (B) root: shoot in the feedback phase on conditioning Cd and plant treatments and feedback species. Means ± 1 s.e. Letters indicate means that did not differ in post hoc tests (P<0.05). Figure 4. The dependence of plant-soil feedback index (PSF= ln (mass in home soil /mass in away soil) on Cd concentrations in the conditioning phase. 95% confidence intervals. Figure 5. Spearman correlations between soil chemical properties, soil microbes, and soil microbes with a mass of P. Americana or P. Acinosa in the feedback phase. Rho and P values. Figure S1. The dependence of plant (A) height, (B) aboveground mass, (C) belowground mass, and (D) root: shoot on soil Cd concentrations and plant species in the conditioning phase. Means ± 1 s.e. Letters indicate means that did not differ in post hoc tests (P<0.05). Figure S2. The dependence of (A) soil bacteria abundance, (B) fungi: bacteria, (C) gram-positive bacteria to gram-negative bacteria, (D) soil water content on soil Cd concentrations and plant treatment in the conditioning phase. Means ± 1 s.e. Letters indicate means that did not differ in post hoc tests (P<0.05). Figure S3. The dependence of plant (A) aboveground mass and (B) belowground mass in the feedback phase on conditioning Cd and plant treatments and feedback species. Means ± 1 s.e. Figure S4. Spearman correlations between soil microbes and soil microbes with P. Americana or P mass . Acinosa in the feedback phase. Rho and P values. Supplementary Material File (image1.emf) Download 60.61 KB File (image2.emf) Download 91.42 KB File (image3.emf) Download 76.59 KB File (image4.emf) Download 17.54 KB File (image5.emf) Download 22.70 KB File (image6.emf) Download 60.16 KB File (image7.emf) Download 61.91 KB File (image8.emf) Download 64.13 KB File (image9.emf) Download 16.65 KB File (oik-11452-file003.docx) Download 18.28 KB File (oik-11452-file004.docx) Download 83.49 KB References 1. Bates SE, Wandrag EM, Duncan RP (2020). Calculating the uncertainty associated with log response ratios in plant–soil feedback studies. Plant Ecol 221: pp. 829–836. 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Crossref Google Scholar Information & Authors Information Version history V1 Version 1 21 February 2025 Peer review timeline Published Journal Of Plant Ecology Version of Record 29 Aug 2025 Published Copyright This work is licensed under a Non Exclusive No Reuse License. Keywords heavy metal cd phytolacca americana plant invasion plant-soil feedback soil secondary metabolites soil biota Authors Affiliations Yue Zhu 0009-0009-3207-2870 View all articles by this author Jun Chen View all articles by this author Yu Ming View all articles by this author Jingru Zhang View all articles by this author Shaoyu Zhang View all articles by this author Yunshan Liu View all articles by this author Bo Li Yunnan University View all articles by this author Jihua Wu 0000-0001-8623-8519 State Key Laboratory of Herbage Improvement and Grassland Agro-Ecosystems, College of Ecology, Lanzhou, University, Lanzhou, 730000, China View all articles by this author Evan Siemann Rice University View all articles by this author Yi Wang 0000-0003-1851-8994 [email protected] Yunnan University View all articles by this author Metrics & Citations Metrics Article Usage 227 views 147 downloads .FvxKWukQNSOunydq8rnd { width: 100px; } Citations Download citation Yue Zhu, Jun Chen, Yu Ming, et al. Effects of cadmium pollution on soil feedback between invasive plant Phytolacca Americana and native plant Phytolacca Acinosa. Authorea . 21 February 2025. DOI: https://doi.org/10.22541/au.174013593.35371378/v1 If you have the appropriate software installed, you can download article citation data to the citation manager of your choice. Simply select your manager software from the list below and click Download. For more information or tips please see 'Downloading to a citation manager' in the Help menu . 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