Solidago canadensis and its endophytes as a self-degrading system during ex situ decomposition

Solidago canadensis and its endophytes as a self-degrading system during ex situ decomposition | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Solidago canadensis and its endophytes as a self-degrading system during ex situ decomposition Kamil Kisło, Patryk Czortek, Anna Wiewiorowska, Marta Wrzosek This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8917351/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 6 You are reading this latest preprint version Abstract Background and Aims Solidago canadensis is an invasive herb widespread across Eurasia and Australia, known to decompose faster than native species. However, the mechanisms driving its decomposition, particularly the role of associated microorganisms, remain poorly understood. This study aimed to assess the influence of specific components of soil biota (bacteria and fungi) from invaded and non-invaded mesic meadows on the decomposition of S. canadensis . Methods We conducted an ex situ decomposition experiment using shoots and leaves of S. canadensis . Soil solutions from invaded and non-invaded meadows were prepared and passed through syringe filters of varying pore sizes (variants A–D). We assessed primary decomposer diversity under a light microscope and analysed mass loss using generalized linear mixed models. Results Differences among filtration treatments were minimal, and soil solution origin (invaded vs. non-invaded meadows) had negligible effects on mass loss. Leaves and shoots decomposed at similar rates across all variants. These outcomes indicate that external microbial inputs played no detectable role in driving decomposition. Conclusion Our findings suggest that S. canadensis operates as a largely self-contained holobiont, harbouring an internal consortium of primary decomposers capable of sustaining decomposition independently of external soil communities. This highlights the functional redundancy of surrounding soil biota and may contribute to enhanced nutrient cycling in invaded ecosystems. Such a strategy aligns with invasion facilitation frameworks, including the novel weapon hypothesis, illustrating how S. canadensis can exploit new environments. Additionally, its consistently efficient ex situ decomposition suggests strong potential for valorizing harvested biomass through composting or biogas production. Solidago canadensis invasive plant environmental engineer fungi bacteria decomposition Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Introduction Decomposition is a crucial part of the carbon cycle, depending, among others, on the activity of the microorganisms and mesofauna. Some components of this biota can be directly responsible for mass loss through the use of their enzymes and feeding on detritus (primary decomposers), or indirectly regulate the populations of other organisms (secondary decomposers) (Hättenschwiler et al., 2005 ). Both of these groups contain representatives of fungi, bacteria, protists, and mesofauna. Hättenschwiler et al. ( 2005 ) suggested that bacteria and fungi are the main primary decomposers since they are the key organisms transforming detritus and regulating mesofauna’s activity. Despite the high phylogenetic heterogeneity, all of the components of decomposing biota synergistically interact with each other (i.e., commensalism, mutualism, synergistic decomposition of specific compounds), or antagonistically (i.e., competition, predation, parasitism), and the mechanisms of these interactions may vary depending on abiotic environmental factors, such as light and water availability, temperature, or type of substrate (Ferreira and Chauvet, 2010; Tylianakis et al., 2008 ). Decomposition, as a complex of environmental processes, remains incompletely understood despite a rapidly expanding body of literature. Its dynamics are strongly context-dependent and vary across habitats, biogeographic regions, and spatial scales. In the Anthropocene, biological invasions have emerged as an additional and still insufficiently recognised driver that can substantially alter decomposition processes. Invasive species can disrupt ecosystem functioning, especially when they act as “environmental engineers” (Czortek et al., 2026 ; Xin et al. , 2022). By introducing novel biotic and abiotic components into the system, they contribute to the formation of novel ecosystems. As a result, they increase both functional and phylogenetic novelty (Schittko et al. 2020 ), which may shape decomposition processes in ways that are only beginning to be understood. Environmental engineers can influence soil moisture by modifying river and groundwater dynamics (Stromberg et al ., 2007). They may also alter light penetration (Dyderski and Jagodziński, 2019) and shift soil nutrient processes (Ehrenfeld, 2003). In addition, they can affect litter decomposition (Castro-Díez et al ., 2009, 2019). Their presence may further lead to novel interactions with native biota, as tissues of some invaders decompose at a similar rate across all stages of the invasion gradient (Kisło et al., 2025 ). One of the invasive species belonging to the group of environmental engineers is Solidago canadensis (L.), a member of the Asteraceae family with a native range in North America, and indroduced range in Europe, Asia, and Australia (Ren et al., 2020 ; Zhang et al., 2009 ). It was introduced in Europe relatively recently, with the beginning of its history in the 17th century in England, with further spread in Western and Eastern Europe (Popai and Parker, 2014). Nevertheless, S. canadensis is considered invasive in urban and suburban environments, wetlands, forests, grasslands, and meadows, including mesic meadows protected by the Natura 2000 (Akatov et al., 2021 ; Kotowska et al. 2024 ; Tokarska-Guzik, 2005). Solidago canadensis is known for its engineering abilities, which include altering soil pH and the C:N ratio (Xie et al., 2023 ) and secreting allelopathic secondary compounds (Anžlovar et al., 2020 ; Deng et al., 2015 ). It also modifies the structure and species composition of soil biota. For example, Xu et al. ( 2024 ) showed that the dominant arbuscular mycorrhizal fungi in the rhizosphere shifted from Glomus to Paraglomus, thereby affecting interactions within soil-inhabiting communities. Nevertheless, the knowledge about the impact of these drivers on the effectiveness of primary decomposers is still limited. Regarding the leaf litter of S. canadensis , different research reveals different decomposition speeds, but most of them suggest that S. canadensis leaf litter decomposes faster than selected species of native flora (Simon et al. 2023 ; Zhang et al. 2016 ), demonstrating, however, high context dependency of these results. This context-dependency depends strongly on the ecosystem type and accompanying environmental conditions. For example, Simon et al. ( 2023 ) investigated a lake ecosystem, whereas Zhang et al. ( 2016 ) examined coastal grasslands. According to this context dependency, our knowledge about microorganisms involved in the S. canadensis decomposition process and their interactions is limited. Since most of the decomposition experiments were conducted in situ (Latterini et al. 2023 ; Simon et al. 2023 ; Zhang et al. 2024), the authors were not able to exclude the impact of a complex of factors (i.e., weather, sunlight exposure, moisture, human activity) on the biogeochemical processes. Despite the high utility of the results of these studies, many applied ecology processes (e.g., composting or biogas production) are conducted ex-situ and need other gaps of knowledge to be filled for their optimisation, including microorganisms taking part in this process and even the decomposition speed. We believe that connecting the experimentally identified, normalised decomposition speed ex-situ with the interactions between the decomposers can be highly useful for further research on this topic. The main goal of this study was to evaluate the impact of fungi and bacteria with different origins on S. canadensis shoots and leaves decomposition in the ex situ conditions of constant moisture and temperature. We decided to perform an ex situ experiment to eliminate the influence of weather conditions on decomposition. This approach also allowed us to control for spatial microheterogeneity in soil properties, including moisture, pH, and fertility. Thus, we focused specifically on the effects of different-sized organisms on the decomposition process, with particular emphasis on bacteria and fungi as the primary decomposers. We decided to collect the information about the influence of bacteria and fungi on S. canadensis leaves and shoots decomposition by eliminating specific components of the soil biota from the controlled experimental setup and comparing the results before and after the removal. We hypothesized that fungi and bacteria would have similar effects on the decomposition (H1) . We hypothesize that representatives of both microorganisms’ kingdoms will act in parallel in the process of decomposition; both bacteria and fungi possess a broad pool of enzymes and can degrade hard organic material (eg, lignin; Bugg et al., 2011 ). Moreover, we expect them to have a positive impact on the decomposition, since Kisło et al. ( 2025 ) suggested that the taxonomic diversity of fungi was positively correlated with the mass loss. On the other hand, if there is only a slight difference between the experimental variants could show us that S. canadensis poses a higher than previously known threat to biodiversity since microbiota living on or inside the plant can decompose its tissues without any additional biotic components. Materials and methods In October 2023, we collected senescing leaves and shoots of S. canadensis and dried them to constant mass (60℃ for 48 hours). During this period, we also collected 100 g of soil from five plots with high S. canadensis cover (ranging from 85% to 95%) and control plots (with less than 5% S. canadensis cover) located in mesic grasslands in Northeastern Poland (Central-Western Europe). The place of soil collection in the current manuscript is referred to as “soil origin”, and the coordinates of these study plots are available in Supplementary Material S1. Before starting the experiment, we dried the material at 70°C for 72 hours to exclude as much epiphytic biota as possible. We prepared mesh bags containing 5g of the material +/- 2% per each meshbag (leaves or shoots, mesh size 1mm) and soil solution with previously sampled soil and 200 ml of autoclaved distilled water. To check the impact of different fractions of soil biota on the decomposition, we prepared four variants of the experiment in sterile plastic containers. Experimental variants were as follows: A − 30 mL of unfiltered soil solution (with all of the biota fraction present), B − 30 mL of soil solution filtered through a 10 µm syringe filter, which excluded mesofauna and large protists from the soil solution, C − 30 mL of soil solution filtered through a 1.2 µm syringe filter, which excluded most of the biota from the soil solution. After that, we added 270 mL of autoclaved distilled water to each container. To check the effect of remaining endophytic biota on the decomposition, we prepared the control variant (D) with 300 mL of the autoclaved distilled water. After the preparation of all experimental variants, we put four meshbags containing leaves or shoots into each of them (containers with leaves, total n = 160, containers with shoots, total n = 160) (Fig. 1 ). Containers were incubated at room temperature. We performed sampling four times per one S. canadensis organ variant: once every six weeks for leaves and once every twelve weeks for shoots, since shoots decompose approximately 50% slower (Anda et al., 2023 ). The remaining necromass from each meshbag was dried and weighed. To obtain as much biodiversity data as possible, during each sampling event, we collected five ml of suspension in which the material was immersed and five mL of water from squeezing the meshbag. These solutions were combined, and 1.5 ml of the mixture was transferred into a sterile Eppendorf tube. After centrifugation (5000 rpm, 5 min), we discarded the supernatant and resuspended the pellets in 100 µl of distilled water. We observed 25 µl of this sample under the light microscope (Nikon Eclipse Ni-U) with a gridded cover slide to count the number of bacterial cells within one field of view, and animals, fungi, and protists within 50 fields of view (microscope magnification x200). We identified the representatives of the last two groups to the genus level (Seifert et al. , 2011; Watanabe, 2002 ) and counted them. To calculate the number of bacterial cells, we placed a 100µm x 100µm square at a randomly chosen point in one randomly chosen field of view from each plot on each collection date. Despite the detection of animals and protists in several samples (Supplementary Material S2), we have decided to focus on the influence of the primary decomposers on the mass loss. To obtain the information about mass loss, we dried the material in the meshbag as at the beginning of the experiment and weighed to assess the mass loss within one sample. Statistical analysis We performed statistical analysis in the R Studio version. 4.3.2 (R Core Team 2025 ). In all of the models response variable was defined by us as a proportion of remaining mass within the sample to the initial mass (5g), hereinafter referred to as “mass loss”. We calculated taxonomic richness of fungi using the vegan::specnumber () function (Oksanen et al., 2025 ). We performed two-stage statistical analyses to obtain information about the influence of the fungal richness and bacterial abundance on the mass loss. At the beginning, we calculated the generalised linear mixed models using glmmTMB:glmmTMB() function (Brooks et al., 2017 ), and then we calculated marginal responses of each model using ggeffects :: ggpredict() function (Lüdecke, 2016) and marginal means using emmean::emmeans() function (Lenth, 2023) to model the influence of single predictor, handling other predictors at a constant (mean) level. We used the ggplot2 :: ggplot () function (Wickham, 2009 ) to visualise the marginal responses. All of the models contained a box ID (eg, S33D) nested in plot ID (eg, S33) as a random intercept. For variant D (only distilled water added), we defined “plot ID” and “soil origin” as “none”. Fragments of dark septated mycelium (DSM) and hyaline mycelium can serve as propagules in some fungal taxa and play different ecological roles (Seifert et al. , 2011). We decided to calculate their abundance. To perform this analysis, we used the difference between the number of hyaline mycelium fragments and the number of melanised mycelium fragments (H-DR) as a predictor. Models for leaf litter decomposition were as follows: Eq. 1 = mass loss ~ experimental variant+soil origin+collection+experimental variant:soil origin+experimental variant:collection+collection:soil origin Eq. 2 = mass loss ~ fungal taxonomic richness + fungal taxonomic richness:experimental variant + fungal taxonomic richness:soil origin + fungal taxonomic richness:collection Eq. 3 = mass loss ~ H-DR + H-DR:experimental variant + H-DR:soil origin + H-DR:collection Eq. 4 = mass loss ~ bacteria relative abundance + bacteria relative abundance:experimental variant + bacteria relative abundance:soil origin + bacteria relative abundance:collection. Models for shoots decomposition were as follows: Eq. 5 = experimental variant+soil origin+collection+experimental variant:soil origin+experimental variant:collection+collection:soil origin Eq. 6 = mass loss ~ fungal taxonomic richness + fungal taxonomic richness:experimental variant + fungal taxonomic richness:soil origin + fungal taxonomic richness:collection Eq. 7 = mass loss ~ H-DR + H-DR:experimental variant + H-DR:soil origin + H-DR:collection Eq. 8 = mass loss ~ bacteria relative abundance + bacteria relative abundance:experimental variant + bacteria relative abundance:soil origin + bacteria relative abundance:collection . Results We identified 25 different fungal morphotypes. The most common morphotypes were Fusarium sp., Penicillium sp./ Aspergillus sp., and Cladosporium sp., and the mean spores per sample was 31.08 with 40.38 SD. The mean number of bacterial cells per 100µm x 100µm square was 77.41 with 62.09 SD. Leaves decomposition After the first six weeks of the experiment, the mean mass loss was 34.67% (± 2.40% SD). After 12 weeks, it was 45.55% (± 3.41% SD), after 18 weeks, 49.78% (± 1.77% SD), and after 24 weeks, 52.31% (± 3.59% SD). Mean mass loss at the end of experiment was 52.82% (± 2.43% SD) in variant A (without filtering), 54.08% (± 3.86% SD) in variant B with filtering through a 10um filter, 52.04% (± 4.39% SD) in variant C with filtering through a 1.2um filter, and 53.64% (± 2.37% SD) in control (Fig. 2 ; Eq. 1). Overall, fungal taxonomic diversity was responsible for a 10% mass loss increase (from 43% with one morphotype detected to 51% with 12 morphotypes). The addition of soil solution from invaded plots and control plots in interaction with fungal richness was responsible for 12% and 10% mass loss, respectively. After six weeks of the experiment, the fungal taxonomic diversity was responsible for a 14% decrease in the mass loss, and after 24 weeks for a 16% increase. In variant A, fungal diversity was responsible for 7% mass loss decrease, in variant B for a decrease, in variant C for 7% decrease, and in variant D for 2% increase (Fig. 3 ; Eq. 2). When the abundance of DSM was higher than the abundance of hyaline mycelium decomposition ratio was also higher. Mass loss was 56% when the H-DR (hyaline minus dark ratio) was 10, and when the H-DR was 18, the mass loss ratio was 30%. There was no difference in the influence of mass loss between different soil origins in the case of the H-DR. After six weeks, the higher abundance of hyaline mycelium was responsible for a 21% decrease in the mass loss, and after 24 weeks, for a 13% decrease. In variant A, the higher abundance of hyaline mycelium was responsible for a 26% mass loss decrease, in variant B and D for a 27% mass loss decrease, and in variant C for an 18% mass loss decrease (Fig. 4 ; Eq. 3). Overall, bacterial relative abundance was responsible for 3% mass loss increase (from 48% with 20% of bacteria to 50% with 100% relative bacterial abundance). Bacterial relative abundance effects on the mass loss varied from negligible to weak (2% − 8%) in the interaction with other studied parameters. (Fig. 5 ;Eq. 4). Shoots decomposition Decomposition of shoots was slower than that of leaves, even with longer periods between collections. After the first 12 weeks of the experiment, the mean mass loss was 15.41% (± 2.36% SD), after 24 weeks, 18.45% (± 5.74% SD), after 36 weeks, 28.67% (± 7.39% SD), and after 48 weeks 30.66% (± 7.87% SD). Mean mass loss at the end of an experiment was 30.66% (± 7.87% SD) in variant A (without filtering), 54.08% (± 3.86% SD) in variant B with filtering through a 10um filter, 52.04% (± 4.39% SD) in variant C with filtering through 1.2um filter, and 53.64% (± 2.37% SD) in control (Fig. 2 ; Eq. 5). Overall, fungal taxonomic richness was responsible for a 7% mass loss decrease (from 21% with one morphotype detected to 15% with 10 morphotypes). The addition of soil solution from invaded plots and control plots in interaction with fungal richness was responsible for a 6% and 5% mass loss decrease, respectively. After 12 weeks of the experiment, fungal taxonomic richness was responsible for a 14% decrease in the mass loss, and after 48 weeks for 18% increase. In variant A and C, fungal richness was responsible for 7% mass loss decrease, in variant B for a 10% decrease, and in variant D for 2% decrease (Fig. 3 ;Eq. 6). When the abundance of dark septated mycelium (DSM) was higher than the abundance of hyaline mycelium, the decomposition ratio was also higher. Mass loss was 29% when the H-DR (hyaline minus dark ratio) was − 5, and when the H-DR was 20, the mass loss ratio was 9%. Higher abundance of hyaline mycelium was responsible for a 25% decrease in the decomposition in variants with invasive soil added, and a 20% decrease in variants with native soil added. After 12 weeks, the higher abundance of hyaline mycelium was responsible for a 21% decrease in the mass loss, and after 48 weeks for a 13% decrease. In variant A, the higher abundance of hyaline mycelium was responsible for a 21% mass loss decrease, in variant B for a 10% mass loss decrease, and in variants C and D for a 20% mass loss decrease (Fig. 4 ; Eq. 2). Overall, bacterial relative abundance was responsible for a 12% mass loss decrease (from 22% with 20% of bacteria to 12% with 100% relative bacterial abundance). Soil origin had a negligible effect on the influence of the bacterial abundance on the mass loss. After 12 weeks of the experiment, bacterial relative abundance was responsible for a 13% decrease in the mass loss, and after 48 weeks for 7% increase. In all experimental variants, the abundance of bacteria had a similar negative effect (10% to 14% mass loss decrease, Fig. 5 ; Eq. 5). Discussion Mass loss within different experimental variants In the current study, the mass loss grew faster than in earlier studies on S. canadensis litter decay conducted in the terrestrial environments (e.g., Zhang et al., 2016 ). Nevertheless, the mass loss we obtained was close to the S. canadensis shoot and leaf decomposition ratios reported by Anda et al. ( 2023 ) in the freshwater. This similarity was expected, as both our experiment and that of Anda et al. eliminated the negative effects of drought on microbial activity through their experimental setups. On the other hand, we expected more factors in our study to affect the decomposition. First of all, our expectations regarding the impact of soil origin (invaded vs. control plots) were not confirmed. We observed only negligible differences between filtration variants of soil solution and variants of suspended soil (from Solidago stands, and mesic meadows without Solidago ). Moreover, our data do not show any consistent trend in these relationships. For instance, after 36 weeks, variants with soil from non-invaded plots had slightly higher mass loss than those with soil solution from invaded ones, but after another 12 weeks, their effects were relatively similar. As shown by Kisło et al. ( 2025 ), the increasing cover of S. canadensis on experimental plots does not affect the rate of leaf litter decomposition. Moreover, the main part of this study - different fractions of soil solutions expressed as soil solutions’ filtering variants - turned out to be ineffective predictors of S. canadensis litter decomposition as soil solution origin (i.e., invaded and non-invaded meadows). As regards soil origin, we observed slight differences between the decomposition speed within different filtration variants, but they are hard to interpret ecologically. They rather derive from interspecies litter quality (e.g., organs’ thickness, or lignin concentration), which may exert a pronounced impact on the decomposition (Couˆteaux et al. , 1995), than from ecological processes concerning the impact of saprotrophic biota. Moreover, this indicates a high degree of functional redundancy among S. canadensis decomposers. More specifically, when one functional group was removed or underperformed, others were able to compensate and maintain decomposition processes. For instance, the elimination of soil-borne fungi allowed bacteria to occupy more environmental niches, thus degrading more types of substrates, since the difference in mass loss between variants A, B, and C was negligible. Moreover, in our samples, we observed Harzia sp., a mycopathogenic fungus (Nguyen et al., 2016 ), which may have regulated the fungal population, and by that, the fungal impact on the decomposition, when other fungivorous organisms were missing. Variant D (control), consisting solely of S. canadensis leaves/shoots and their native biota, showed a mass loss level comparable to the other variants. This allows for a cautious hypothesis that the microbiota naturally inhabiting S. canadensis tissues may contribute to the decomposition of its own litter. Endophyte-driven priming has already been demonstrated (Guerreiro et al. , 2017), and our results may indicate a similar but not yet fully confirmed mechanism. Placed in the hologenome context (Moran and Sloan, 2015 ), this would mean that the plant and its tissue-associated microbes form a unit capable of recycling nutrients embedded in the biomass. This link is particularly relevant for invasive species such as S. canadensis , which have a high nutrient demand due to their reproduction strategy based on excessive diaspore production (Hua et al. , 2007; Ye et al. , 2018). Moreover, S. canadensis represents a “try harder” strategy, characterized by an acquisitive pattern of resource use aimed at maximizing growth and reproductive output (Tecco et al., 2010 ). Because these nutrients are strongly accumulated in the leaves, conditions are favourable for the development of saprotrophs within the plant’s own tissues. This may explain both the similarity between our results and those of Anda et al. ( 2023 ) and the limited effect of soil origin or filtering. S. canadensis may already carry saprotrophs capable of initiating decomposition, supporting nutrient return to the plant or its offspring. Bacteria and fungi have similar effects on the decomposition Unexpectedly, the relative abundance of bacteria and fungal diversity showed a negative correlation with mass loss at the beginning of the experiment. Plant litter with S. canadensis origin may be hard to degrade due to high lignin concentration in shoots (Wiatrowska et al. 2022 ). Probably cuticle, as the natural barrier between inner plant tissues and the environment, needed the initial degradation process, as shown in (Logan et al. , 2022). Moreover, S. canadensis is known for producing a high diversity of secondary metabolites with different stability and potential impact on the decomposition (Zaimenko et al., 2025 ). Nevertheless, a high abundance of microorganisms that are not rapid decomposers may contribute to the residual mass, thereby slowing the overall rate of mass loss. The production of secondary metabolites in the cuticule-related tissues in S. canadensis organs may be the reason for decreasing the impact of fungi and bacteria on the mass loss at the beginning of the experiment in both shoots and leaves. Notable inhibition by these groups’ representatives was visible after six weeks of experiment in the case of leaf litter and after 12 and 24 weeks in the case of shoots, and we found a probable explanation for this phenomenon. Secondary metabolites of S. canadensis may differ between shoots and leaves, or among different organs. Anžlovar et al. ( 2020 ) showed this experimentally: extracts from different organs had varying effects on Botrytis cinerea growth. Studies by Reyes-Ávila et al. ( 2024 ) and Jenner et al. ( 2011 ) have shown that two of the secondary metabolites present in S. canadensis tissues, germacene D and limonene, have different decomposition ratios. Limonene decomposes fast, in up to seven days in the soil, and up to 40 days in the distilled water (Reyes-Ávila et al. 2024 ), while more than 75% of germacene D survives 63 days of biodegradation (Jenner et al. 2011 ). Moreover, we can expect that the substrate itself remains largely unavailable to saprotrophs, yet the aquatic environment becomes sufficiently enriched with dissolved organic compounds to stimulate bacterial cell division. Thus, bacterial abundance may increase even though the substrate is not yet undergoing intensive decomposition. Since the current study focuses mainly on the fungal idiophase (spores), and sporulation sometimes requires specific conditions (Morton, 1961 ), we also checked the impact of the trophophase (vegetative mycelium) on the decomposition. Despite the knowledge that fungi with a trophophorm of hyaline mycelium have fast metabolism and are effective degraders (i.e., Basidiomycetes (Pleurotus), Mucoromycetes; Cohen et al. , 2002; Satari and Karimi, 2017), our results suggest the opposite dependence. Some plant-associated fungi with melanised hyphae, such as Cladosporium spp., show broad metabolic and functional diversity (Bensch et al., 2018 ; Nguyen et al., 2016 ). However, this factor alone does not explain why our findings differ from those of the field study by Kisło et al. ( 2025 ), in which DSM had a clearly negative effect on mass loss. One possible explanation is that under constant high humidity and optimal temperature, DSM fungi may allocate more resources to faster trophic metabolism. Another possibility is that environmental conditions select for different taxonomic groups of DSM-associated decomposers in different settings. Unfortunately, our experiment could not determine which of these two possibilities is closer to the truth. Taken together, these patterns indicate that bacteria and fungi exert broadly similar effects on the decomposition of both leaves and shoots, regardless of soil solution origin. This consistency suggests that external microbial communities play only a minor role in shaping early and mid-term decomposition dynamics in S. canadensis . Study limitations Our experiment was conducted at room temperature, which removed the effect of varying environmental conditions but may have limited the activity of fungi requiring stratification for germination (Juge et al. , 2002). We also detected fungal spores and nematodes in soil-filtration variants. Because their presence on leaves and shoots was more likely than filtration errors, we treated them as components of the endophytic/epiphytic biota of S. canadensis . Only direct microscopy was used, as culturing would require partitioning the material and could introduce decomposition-rate biases. For the same reason, we excluded the metabarcoding approach. Using soil for experimental preparation also increased the risk of detecting dormant spores or eggs unrelated to S. canadensis decomposition, potentially producing false metabarcoding signals. Moreover, the magnification used allows for a relative comparison of bacterial abundance in the sample, but without specific staining, some fraction of the bacteria may be underestimated. Moreover, some bacteria occur in colonies, and such colonies are likely to be analysed as single units. Future experiments should therefore include two storage variants-room temperature and conditions matching the litter collection site-to avoid limiting fungal stratification. Metabarcoding should be performed before the experiment (on fresh and dried litter) to confirm the presence or absence of non-endophytic biota. Additionally, using litter from plants grown under sterile conditions would help eliminate biases introduced by endophytic communities. Conclusions Solidago canadensis is known for acting as an environmental engineer and changing the properties of the environment in the introduced range. When we add novel interactions with endophytic biota acting as efficient decomposers in the introduced range, as stated in Kisło et al. ( 2025 ) and in the current study, we obtain a view of a self-sufficient invader, and by that, fully adapted to modifying new areas. Our concern shall be especially important for lawmakers preparing local lists of invasive species with the highest impact on the environment and the lists of non-native species that should be eradicated from their secondary range. The second, more optimistic view of our results provides a valuable insight into the applied usage of S. canadensis . Since its tissues contain biota that are sufficient decomposers, the effort put into the composting of this plant seems to be relatively low, thus with a relatively low cost. In this case, our results may be part of the opportunity to fight the invasion of this plant, since mowing its shoots twice a year, according to the instructions (Gala-Czekaj et al., 2021 ), may be profitable for both farmers and companies selling bioproducts. Declarations Acknowledgments Author Contributions All authors contributed to the conception and design of the study. Kamil Kisło, Patryk Czortek, Anna Wiewiorowska, and Marta Wrzosek performed material preparation, data collection, and analysis. The first draft of the manuscript was written by Kamil Kisło, and all authors commented on previous versions. All authors read and approved the final manuscript. Data Availability The datasets generated and/or analyzed during the current study are available from the corresponding author upon reasonable request. 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Mycorrhiza 12:37–42. https://doi.org/10.1007/s00572-001-0151-8 Kisło K, Czortek P, Mazurkiewicz MT et al (2025) Breaking down the invader of mesic meadows: fungi accelerate Solidago canadensis leaf litter decay more than protists at the first six months of decomposition. Biol Invasions 27:219. https://doi.org/10.1007/s10530-025-03673-7 Kotowska D, Skórka P, Pärt T et al (2024) Spatial scale matters for predicting plant invasions along roads. J Ecol 112:305–318. https://doi.org/10.1111/1365-2745.14234 Latterini F, Dyderski MK, Horodecki P et al (2023) The Effects of Forest Operations and Silvicultural Treatments on Litter Decomposition Rate: a Meta-analysis. Curr Forestry Rep 9:276–290. https://doi.org/10.1007/s40725-023-00190-5 Lüdecke D (2018) ggeffects: Tidy Data Frames of Marginal Effects from Regression Models. J Open Source Softw 3:772. https://doi.org/10.21105/joss.00772 Moran NA, Sloan DB (2015) The Hologenome Concept: Helpful or Hollow? PLoS Biol 13:e1002311. https://doi.org/10.1371/journal.pbio.1002311 Morton AG (1961) The induction of sporulation in mould fungi. Proc R Soc Lond B Biol Sci 153:548–569. https://doi.org/10.1098/rspb.1961.0018 Nguyen NH, Song Z, Bates ST et al (2016) FUNGuild: An open annotation tool for parsing fungal community datasets by ecological guild. Fungal Ecol 20:241–248. https://doi.org/10.1016/j.funeco.2015.06.006 Oksanen J, Simpson G, Blanchet F, Kindt R, Legendre P, Minchin P, O'Hara R, Solymos P, Stevens M, Szoecs E, Wagner H, Barbour M, Bedward M, Bolker B, Borcard D, Borman T, Carvalho G, Chirico M, De Caceres M, Durand S, Evangelista H, FitzJohn R, Friendly M, Furneaux B, Hannigan G, Hill M, Lahti L, Martino C, McGlinn D, Ouellette M, Ribeiro Cunha E, Smith T, Stier A, TerBraak C, Weedon J (2025) vegan: Community Ecology Package. R package version 2.8-0. https://vegandevs.github.io/vegan/ Popai I, Parker C (2019) Solidago canadensis (Canadian goldenrod). CABI Compendium Ren G, He M, Li G et al (2020) Effects of Solidago canadensis Invasion and Climate Warming on Soil Net N Mineralization. Pol J Environ Stud 29:3285–3294. https://doi.org/10.15244/pjoes/114237 R Core Team (2025) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. https://www.R-project.org/ Reyes-Ávila A, Garrido Frenich A, Romero-González R (2025) Degradation Study of Limonene and trans-Cinnamaldehyde in Water Samples: Sunlight and Darkness Studies. Water Air Soil Pollut 236:306. https://doi.org/10.1007/s11270-025-07932-0 Reyes-Ávila A, Garrido Frenich A, Romero-González R (2024) Degradation of limonene and trans-cinnamaldehyde in soil, and detection of their metabolites by UHPLC and GC-HRMS. Environ Sci Pollut Res 31:33058–33068. https://doi.org/10.1007/s11356-024-33334-6 Satari B, Karimi K (2018) Mucoralean fungi for sustainable production of bioethanol and biologically active molecules. Appl Microbiol Biotechnol 102:1097–1117. https://doi.org/10.1007/s00253-017-8691-9 Schittko C, Bernard-Verdier M, Heger T et al (2020) A multidimensional framework for measuring biotic novelty: How novel is a community? Glob Chang Biol 26:4401–4417. https://doi.org/10.1111/gcb.15140 Schneider T, Keiblinger KM, Schmid E et al (2012) Who is who in litter decomposition? Metaproteomics reveals major microbial players and their biogeochemical functions. ISME J 6:1749–1762. https://doi.org/10.1038/ismej.2012.11 Seifert KA, Gams W (2011) The genera of Hyphomycetes – 2011 update. Persoonia - Mol Phylogeny Evol Fungi 27:119–129. https://doi.org/10.3767/003158511X617435 Simon S, Simon-Gáspár B, Anda A (2023) Carbon dioxide emission and its impacting factor from goldenrod (Solidago canadensis) and bulrush (Typha angustifolia) decomposition during the winter period. Ecohydrol Hydrobiol 23:177–185. https://doi.org/10.1016/j.ecohyd.2023.01.002 Tecco PA, Díaz S, Cabido M, Urcelay C (2010) Functional traits of alien plants across contrasting climatic and land-use regimes: do aliens join the locals or try harder than them? J Ecol 98:17–27. https://doi.org/10.1111/j.1365-2745.2009.01592.x Tokarska - Guzik B (2005) The establishment and spread of alien plant species (Kenophytes) in the flora of Poland. Wydawnictwo Uniwersytetu Śląskiego, Katowice Tylianakis JM, Didham RK, Bascompte J, Wardle DA (2008) Global change and species interactions in terrestrial ecosystems. Ecol Lett 11:1351–1363. https://doi.org/10.1111/j.1461-0248.2008.01250.x Watanabe T (2002) Pictorial Atlas of Soil and Seed Fungi. CRC Wiatrowska BM, Wawro A, Gieparda W, Waliszewska B (2022) Bioethanol Production Potential and Other Biomass Energy Properties of Invasive Reynoutria, Solidago, and Spiraea Plants. Forests 13:1582. https://doi.org/10.3390/f13101582 Wickham H (2009) ggplot2. Springer New York, New York, NY Xie H, Knapp LSP, Yu M, Wang GG (2023) Solidago canadensis invasion destabilizes the understory plant community and soil properties of coastal shelterbelt forests of subtropical China. Plant Soil 484:65–77. https://doi.org/10.1007/s11104-022-05739-0 Xu W, Pan Q, Zhang Q et al (2024) Synergistic changes in AM fungi and soil abiotic properties in rhizosphere soils of invasive Solidago canadensis may confer its stronger dominance in communities. Plant Soil. https://doi.org/10.1007/s11104-023-06454-0 Ye X-Q, Yan Y-N, Wu M, Yu F (2019) High Capacity of Nutrient Accumulation by Invasive Solidago canadensis in a Coastal Grassland. Front Plant Sci 10. https://doi.org/10.3389/fpls.2019.00575 Zaimenko N, Chernikova N, Didyk N et al (2025) Phenological changes in secondary metabolites and mineral nutrition of Solidago canadensis and their impact on the rhizosphere soil ecosystem. Plant Introduction 107:36–47. https://doi.org/10.46341/PI2025007 Zhang CB, Wang J, Qian BY, Li WH (2009) Effects of the invader Solidago canadensis on soil properties. Appl Soil Ecol 43:163–169. https://doi.org/10.1016/j.apsoil.2009.07.001 Zhang L, Ma X, Wang H et al (2016) Soil Respiration and Litter Decomposition Increased Following Perennial Forb Invasion into an Annual Grassland. Pedosphere 26:567–576. https://doi.org/10.1016/S1002-0160(15)60066-2 Supplementary Files SupplementarymaterialS1.xlsx supplementarymaterials2.xlsx SupplementarymaterialS3.xlsx Cite Share Download PDF Status: Under Review Version 1 posted Editorial decision: Major revisions 04 May, 2026 Reviewers agreed at journal 26 Feb, 2026 Reviewers invited by journal 26 Feb, 2026 Editor invited by journal 20 Feb, 2026 Editor assigned by journal 19 Feb, 2026 First submitted to journal 19 Feb, 2026 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. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-8917351","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":597375939,"identity":"aa7c88ee-0885-400b-b9a2-feb05ba2aad2","order_by":0,"name":"Kamil Kisło","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAAyUlEQVRIiWNgGAWjYBACCQY2hgMMBQxyYB4PEPMRp8WAwRiuhY0YLQxALYkNRGuRbGBLPPjDwCZ9w40Exgdv2xjyCGqRZmA7cJjHIC0XqIXZcG4bQzFBLXIM7A2HGQwO5264ncAmzdvGkNhGjBagww6nG9xOYP9NlBaQww7wGBxOAGphYyZKi2QzWwLIL4Yz7z9slpxzToKwXySOtxl//FFhI8935vDBD2/KbPL4CWlhYIazGBtARiQQ1IEByNAyCkbBKBgFwx0AANeLOT4fvjZlAAAAAElFTkSuQmCC","orcid":"https://orcid.org/0000-0002-7569-0072","institution":"Botanic Garden, Faculty of Biology, University of Warsaw","correspondingAuthor":true,"prefix":"","firstName":"Kamil","middleName":"","lastName":"Kisło","suffix":""},{"id":597375940,"identity":"3bb1b769-8e22-4068-8767-e5161309c84a","order_by":1,"name":"Patryk Czortek","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Patryk","middleName":"","lastName":"Czortek","suffix":""},{"id":597375941,"identity":"cf3c9f41-9838-433c-b4fb-d2e7515ab01c","order_by":2,"name":"Anna Wiewiorowska","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Anna","middleName":"","lastName":"Wiewiorowska","suffix":""},{"id":597375942,"identity":"0a4f998f-7187-4730-982c-916febeb4a74","order_by":3,"name":"Marta Wrzosek","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Marta","middleName":"","lastName":"Wrzosek","suffix":""}],"badges":[],"createdAt":"2026-02-19 11:56:01","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-8917351/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-8917351/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":103766102,"identity":"a6fad223-62af-4e69-9c06-d2201959fb07","added_by":"auto","created_at":"2026-03-02 16:12:13","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":174405,"visible":true,"origin":"","legend":"\u003cp\u003eExperimental setup scheme. Letters in bold indicate the experimental variant (A-soil solution without filtering, B-soil solution filtered through 10μm syringe filter, C-soil solution filtered through a syringe filter with 1.2 μm pores, D-control variant with destilled water.\u003c/p\u003e","description":"","filename":"Fig.1.png","url":"https://assets-eu.researchsquare.com/files/rs-8917351/v1/9009cf1bb0be8f9f2dae2219.png"},{"id":103765844,"identity":"5f922278-20bb-46f6-b8aa-5acf61da1d55","added_by":"auto","created_at":"2026-03-02 16:10:09","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":275217,"visible":true,"origin":"","legend":"\u003cp\u003eThe influence of experimental variant and soil origin on the mass loss within different collection periods in the following parts of the experiment: a - leaves variant A, b - leaves variant B, c - leaves variant C, d - leaves variant D, e - shoots variant A, f - shoots variant B, g - shoots variant C, h - shoots variant h. Uppercase letters above each panel means: A - variant A (without filtration), B - filtration through 10 μM syringe filter, C - filtration through 1.2 μM syringe filter, D - distilled water.\u003c/p\u003e","description":"","filename":"fig.2.png","url":"https://assets-eu.researchsquare.com/files/rs-8917351/v1/69c53af2aada30dcab880705.png"},{"id":103766084,"identity":"b3694e47-fa5e-4dd6-b951-9c848a5393d0","added_by":"auto","created_at":"2026-03-02 16:12:07","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":501208,"visible":true,"origin":"","legend":"\u003cp\u003eThe influence of fungal taxonomic richness on the mass loss of leaves (a) and shoots (b), and the influence of fungal taxonomic richness in the interaction with soil origin in leaves c) and shoots (d), the influence of the fungal taxonomic richness during different collection periods in leaf (e) and shoots (f) part of the experiment, the influence of fungal taxonomic richness on the mass loss within different experimental variants in the leaves (g) and shoots (h) part of the experiment).\u003c/p\u003e","description":"","filename":"fig.3.png","url":"https://assets-eu.researchsquare.com/files/rs-8917351/v1/d96b56b1a221985553f03e6e.png"},{"id":103765850,"identity":"b749e180-7b2c-4729-a9a3-ffefdc3583a7","added_by":"auto","created_at":"2026-03-02 16:10:18","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":2641828,"visible":true,"origin":"","legend":"\u003cp\u003eThe influence of the difference between the number of hyaline mycelium fragments and dark septate mycelium fragments (H-DR) on the mass loss of leaves (a) and shoots (b), and the influence of H-DR in the interaction with soil origin in leaves c) and shoots (d), the influence of H-DR during different collection periods in leaf (e) and shoots (f) part of the experiment, the influence of H-DR on the mass loss within different experimental variants in the leaves (g) and shoots (h) part of the experiment.\u003c/p\u003e","description":"","filename":"fig.5.png","url":"https://assets-eu.researchsquare.com/files/rs-8917351/v1/7c7ffecfd33faf1c9b641f26.png"},{"id":103765943,"identity":"5168fcf9-d26b-4d5e-adf0-c593621ab560","added_by":"auto","created_at":"2026-03-02 16:11:20","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":515293,"visible":true,"origin":"","legend":"\u003cp\u003eThe influence of the bacterial relative abundance on the mass loss of leaves (a) and shoots (b), and the influence of bacterial relative abundance in the interaction with soil origin in leaves c) and shoots (d), the influence of bacterial relative abundance during different collection periods in leaf (e) and shoots (f) part of the experiment, the influence of bacterial relative abundance on the mass loss within different experimental variants in the leaves (g) and shoots (h) part of the experiment.\u003c/p\u003e","description":"","filename":"fig.4.png","url":"https://assets-eu.researchsquare.com/files/rs-8917351/v1/a22fdae20f8f35ebdc4c4582.png"},{"id":103766368,"identity":"3b6b5aa1-543f-4ac2-8e0c-fa306928dace","added_by":"auto","created_at":"2026-03-02 16:14:13","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":4551148,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8917351/v1/385cdfb1-a9ee-4f6a-b20f-7cdd6a9d1b1d.pdf"},{"id":103765834,"identity":"c604b9ef-8f31-49d0-b2cb-976ff8ead2ba","added_by":"auto","created_at":"2026-03-02 16:09:52","extension":"xlsx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":8960,"visible":true,"origin":"","legend":"","description":"","filename":"SupplementarymaterialS1.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-8917351/v1/945cd6d1f4538a4a6525e057.xlsx"},{"id":103765840,"identity":"6a11a5c4-5b7b-4495-9e51-699a18d4ad2f","added_by":"auto","created_at":"2026-03-02 16:10:07","extension":"xlsx","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":39788,"visible":true,"origin":"","legend":"","description":"","filename":"supplementarymaterials2.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-8917351/v1/bed611dcdabef09ae6be6f23.xlsx"},{"id":103765833,"identity":"9a2dcd3f-03c2-43ef-9da9-33f915ccdacc","added_by":"auto","created_at":"2026-03-02 16:09:50","extension":"xlsx","order_by":3,"title":"","display":"","copyAsset":false,"role":"supplement","size":11007,"visible":true,"origin":"","legend":"","description":"","filename":"SupplementarymaterialS3.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-8917351/v1/d44c039236679fc8b396752b.xlsx"}],"financialInterests":"","formattedTitle":"Solidago canadensis and its endophytes as a self-degrading system during ex situ decomposition","fulltext":[{"header":"Introduction","content":"\u003cp\u003eDecomposition is a crucial part of the carbon cycle, depending, among others, on the activity of the microorganisms and mesofauna. Some components of this biota can be directly responsible for mass loss through the use of their enzymes and feeding on detritus (primary decomposers), or indirectly regulate the populations of other organisms (secondary decomposers) (H\u0026auml;ttenschwiler et al., \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). Both of these groups contain representatives of fungi, bacteria, protists, and mesofauna. H\u0026auml;ttenschwiler et al. (\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2005\u003c/span\u003e) suggested that bacteria and fungi are the main primary decomposers since they are the key organisms transforming detritus and regulating mesofauna\u0026rsquo;s activity. Despite the high phylogenetic heterogeneity, all of the components of decomposing biota synergistically interact with each other (i.e., commensalism, mutualism, synergistic decomposition of specific compounds), or antagonistically (i.e., competition, predation, parasitism), and the mechanisms of these interactions may vary depending on abiotic environmental factors, such as light and water availability, temperature, or type of substrate (Ferreira and Chauvet, 2010; Tylianakis et al., \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Decomposition, as a complex of environmental processes, remains incompletely understood despite a rapidly expanding body of literature. Its dynamics are strongly context-dependent and vary across habitats, biogeographic regions, and spatial scales. In the Anthropocene, biological invasions have emerged as an additional and still insufficiently recognised driver that can substantially alter decomposition processes.\u003c/p\u003e \u003cp\u003eInvasive species can disrupt ecosystem functioning, especially when they act as \u0026ldquo;environmental engineers\u0026rdquo; (Czortek et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2026\u003c/span\u003e; Xin \u003cem\u003eet al.\u003c/em\u003e, 2022). By introducing novel biotic and abiotic components into the system, they contribute to the formation of novel ecosystems. As a result, they increase both functional and phylogenetic novelty (Schittko et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), which may shape decomposition processes in ways that are only beginning to be understood. Environmental engineers can influence soil moisture by modifying river and groundwater dynamics (Stromberg \u003cem\u003eet al\u003c/em\u003e., 2007). They may also alter light penetration (Dyderski and Jagodziński, 2019) and shift soil nutrient processes (Ehrenfeld, 2003). In addition, they can affect litter decomposition (Castro-D\u0026iacute;ez \u003cem\u003eet al\u003c/em\u003e., 2009, 2019). Their presence may further lead to novel interactions with native biota, as tissues of some invaders decompose at a similar rate across all stages of the invasion gradient (Kisło et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eOne of the invasive species belonging to the group of environmental engineers is \u003cem\u003eSolidago canadensis\u003c/em\u003e (L.), a member of the Asteraceae family with a native range in North America, and indroduced range in Europe, Asia, and Australia (Ren et al., \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Zhang et al., \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). It was introduced in Europe relatively recently, with the beginning of its history in the 17th century in England, with further spread in Western and Eastern Europe (Popai and Parker, 2014). Nevertheless, \u003cem\u003eS. canadensis\u003c/em\u003e is considered invasive in urban and suburban environments, wetlands, forests, grasslands, and meadows, including mesic meadows protected by the Natura 2000 (Akatov et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Kotowska et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Tokarska-Guzik, 2005). \u003cem\u003eSolidago canadensis\u003c/em\u003e is known for its engineering abilities, which include altering soil pH and the C:N ratio (Xie et al., \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) and secreting allelopathic secondary compounds (Anžlovar et al., \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Deng et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). It also modifies the structure and species composition of soil biota. For example, Xu et al. (\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) showed that the dominant arbuscular mycorrhizal fungi in the rhizosphere shifted from Glomus to Paraglomus, thereby affecting interactions within soil-inhabiting communities. Nevertheless, the knowledge about the impact of these drivers on the effectiveness of primary decomposers is still limited.\u003c/p\u003e \u003cp\u003eRegarding the leaf litter of \u003cem\u003eS. canadensis\u003c/em\u003e, different research reveals different decomposition speeds, but most of them suggest that \u003cem\u003eS. canadensis\u003c/em\u003e leaf litter decomposes faster than selected species of native flora (Simon et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Zhang et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), demonstrating, however, high context dependency of these results. This context-dependency depends strongly on the ecosystem type and accompanying environmental conditions. For example, Simon et al. (\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) investigated a lake ecosystem, whereas Zhang et al. (\u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) examined coastal grasslands. According to this context dependency, our knowledge about microorganisms involved in the \u003cem\u003eS. canadensis\u003c/em\u003e decomposition process and their interactions is limited.\u003c/p\u003e \u003cp\u003eSince most of the decomposition experiments were conducted \u003cem\u003ein situ\u003c/em\u003e (Latterini et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Simon et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Zhang \u003cem\u003eet al.\u003c/em\u003e 2024), the authors were not able to exclude the impact of a complex of factors (i.e., weather, sunlight exposure, moisture, human activity) on the biogeochemical processes. Despite the high utility of the results of these studies, many applied ecology processes (e.g., composting or biogas production) are conducted \u003cem\u003eex-situ\u003c/em\u003e and need other gaps of knowledge to be filled for their optimisation, including microorganisms taking part in this process and even the decomposition speed. We believe that connecting the experimentally identified, normalised decomposition speed \u003cem\u003eex-situ\u003c/em\u003e with the interactions between the decomposers can be highly useful for further research on this topic.\u003c/p\u003e \u003cp\u003eThe main goal of this study was to evaluate the impact of fungi and bacteria with different origins on \u003cem\u003eS. canadensis\u003c/em\u003e shoots and leaves decomposition in the \u003cem\u003eex situ\u003c/em\u003e conditions of constant moisture and temperature. We decided to perform an \u003cem\u003eex situ\u003c/em\u003e experiment to eliminate the influence of weather conditions on decomposition. This approach also allowed us to control for spatial microheterogeneity in soil properties, including moisture, pH, and fertility. Thus, we focused specifically on the effects of different-sized organisms on the decomposition process, with particular emphasis on bacteria and fungi as the primary decomposers. We decided to collect the information about the influence of bacteria and fungi on \u003cem\u003eS. canadensis\u003c/em\u003e leaves and shoots decomposition by eliminating specific components of the soil biota from the controlled experimental setup and comparing the results before and after the removal. We hypothesized that fungi and bacteria would have similar effects on the decomposition \u003cb\u003e(H1)\u003c/b\u003e. We hypothesize that representatives of both microorganisms\u0026rsquo; kingdoms will act in parallel in the process of decomposition; both bacteria and fungi possess a broad pool of enzymes and can degrade hard organic material (eg, lignin; Bugg et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Moreover, we expect them to have a positive impact on the decomposition, since Kisło et al. (\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2025\u003c/span\u003e) suggested that the taxonomic diversity of fungi was positively correlated with the mass loss. On the other hand, if there is only a slight difference between the experimental variants could show us that \u003cem\u003eS. canadensis\u003c/em\u003e poses a higher than previously known threat to biodiversity since microbiota living on or inside the plant can decompose its tissues without any additional biotic components.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cp\u003eIn October 2023, we collected senescing leaves and shoots of \u003cem\u003eS. canadensis\u003c/em\u003e and dried them to constant mass (60℃ for 48 hours). During this period, we also collected 100 g of soil from five plots with high \u003cem\u003eS. canadensis\u003c/em\u003e cover (ranging from 85% to 95%) and control plots (with less than 5% \u003cem\u003eS. canadensis\u003c/em\u003e cover) located in mesic grasslands in Northeastern Poland (Central-Western Europe). The place of soil collection in the current manuscript is referred to as \u0026ldquo;soil origin\u0026rdquo;, and the coordinates of these study plots are available in Supplementary Material S1. Before starting the experiment, we dried the material at 70\u0026deg;C for 72 hours to exclude as much epiphytic biota as possible. We prepared mesh bags containing 5g of the material +/- 2% per each meshbag (leaves or shoots, mesh size 1mm) and soil solution with previously sampled soil and 200 ml of autoclaved distilled water. To check the impact of different fractions of soil biota on the decomposition, we prepared four variants of the experiment in sterile plastic containers. Experimental variants were as follows: A\u0026thinsp;\u0026minus;\u0026thinsp;30 mL of unfiltered soil solution (with all of the biota fraction present), B\u0026thinsp;\u0026minus;\u0026thinsp;30 mL of soil solution filtered through a 10 \u0026micro;m syringe filter, which excluded mesofauna and large protists from the soil solution, C\u0026thinsp;\u0026minus;\u0026thinsp;30 mL of soil solution filtered through a 1.2 \u0026micro;m syringe filter, which excluded most of the biota from the soil solution. After that, we added 270 mL of autoclaved distilled water to each container. To check the effect of remaining endophytic biota on the decomposition, we prepared the control variant (D) with 300 mL of the autoclaved distilled water. After the preparation of all experimental variants, we put four meshbags containing leaves or shoots into each of them (containers with leaves, total n\u0026thinsp;=\u0026thinsp;160, containers with shoots, total n\u0026thinsp;=\u0026thinsp;160) (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Containers were incubated at room temperature.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eWe performed sampling four times per one \u003cem\u003eS. canadensis\u003c/em\u003e organ variant: once every six weeks for leaves and once every twelve weeks for shoots, since shoots decompose approximately 50% slower (Anda et al., \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The remaining necromass from each meshbag was dried and weighed. To obtain as much biodiversity data as possible, during each sampling event, we collected five ml of suspension in which the material was immersed and five mL of water from squeezing the meshbag. These solutions were combined, and 1.5 ml of the mixture was transferred into a sterile Eppendorf tube. After centrifugation (5000 rpm, 5 min), we discarded the supernatant and resuspended the pellets in 100 \u0026micro;l of distilled water. We observed 25 \u0026micro;l of this sample under the light microscope (Nikon Eclipse Ni-U) with a gridded cover slide to count the number of bacterial cells within one field of view, and animals, fungi, and protists within 50 fields of view (microscope magnification x200). We identified the representatives of the last two groups to the genus level (Seifert \u003cem\u003eet al.\u003c/em\u003e, 2011; Watanabe, \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2002\u003c/span\u003e) and counted them. To calculate the number of bacterial cells, we placed a 100\u0026micro;m x 100\u0026micro;m square at a randomly chosen point in one randomly chosen field of view from each plot on each collection date. Despite the detection of animals and protists in several samples (Supplementary Material S2), we have decided to focus on the influence of the primary decomposers on the mass loss. To obtain the information about mass loss, we dried the material in the meshbag as at the beginning of the experiment and weighed to assess the mass loss within one sample.\u003c/p\u003e \u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eStatistical analysis\u003c/h2\u003e \u003cp\u003eWe performed statistical analysis in the R Studio version. 4.3.2 (R Core Team \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). In all of the models response variable was defined by us as a proportion of remaining mass within the sample to the initial mass (5g), hereinafter referred to as \u0026ldquo;mass loss\u0026rdquo;. We calculated taxonomic richness of fungi using the \u003cem\u003evegan::specnumber\u003c/em\u003e() function (Oksanen et al., \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). We performed two-stage statistical analyses to obtain information about the influence of the fungal richness and bacterial abundance on the mass loss. At the beginning, we calculated the generalised linear mixed models using \u003cem\u003eglmmTMB:glmmTMB()\u003c/em\u003e function (Brooks et al., \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), and then we calculated marginal responses of each model using \u003cem\u003eggeffects\u003c/em\u003e::\u003cem\u003eggpredict()\u003c/em\u003e function (L\u0026uuml;decke, 2016) and marginal means using \u003cem\u003eemmean::emmeans()\u003c/em\u003e function (Lenth, 2023) to model the influence of single predictor, handling other predictors at a constant (mean) level. We used the \u003cem\u003eggplot2\u003c/em\u003e::\u003cem\u003eggplot\u003c/em\u003e() function (Wickham, \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2009\u003c/span\u003e) to visualise the marginal responses. All of the models contained a box ID (eg, S33D) nested in plot ID (eg, S33) as a random intercept. For variant D (only distilled water added), we defined \u0026ldquo;plot ID\u0026rdquo; and \u0026ldquo;soil origin\u0026rdquo; as \u0026ldquo;none\u0026rdquo;. Fragments of dark septated mycelium (DSM) and hyaline mycelium can serve as propagules in some fungal taxa and play different ecological roles (Seifert \u003cem\u003eet al.\u003c/em\u003e, 2011). We decided to calculate their abundance. To perform this analysis, we used the difference between the number of hyaline mycelium fragments and the number of melanised mycelium fragments (H-DR) as a predictor. Models for leaf litter decomposition were as follows:\u003c/p\u003e \u003cp\u003eEq.\u0026nbsp;1\u0026thinsp;=\u0026thinsp;mass loss\u0026thinsp;~\u0026thinsp;experimental variant+soil origin+collection+experimental variant:soil origin+experimental variant:collection+collection:soil origin\u003c/p\u003e \u003cp\u003eEq.\u0026nbsp;2\u0026thinsp;=\u0026thinsp;mass loss\u0026thinsp;~\u0026thinsp;fungal taxonomic richness\u0026thinsp;+\u0026thinsp;fungal taxonomic richness:experimental variant\u0026thinsp;+\u0026thinsp;fungal taxonomic richness:soil origin\u0026thinsp;+\u0026thinsp;fungal taxonomic richness:collection\u003c/p\u003e \u003cp\u003eEq.\u0026nbsp;3\u0026thinsp;=\u0026thinsp;mass loss\u0026thinsp;~\u0026thinsp;H-DR\u0026thinsp;+\u0026thinsp;H-DR:experimental variant\u0026thinsp;+\u0026thinsp;H-DR:soil origin\u0026thinsp;+\u0026thinsp;H-DR:collection\u003c/p\u003e \u003cp\u003eEq.\u0026nbsp;4\u0026thinsp;=\u0026thinsp;mass loss\u0026thinsp;~\u0026thinsp;bacteria relative abundance\u0026thinsp;+\u0026thinsp;bacteria relative abundance:experimental variant\u0026thinsp;+\u0026thinsp;bacteria relative abundance:soil origin\u0026thinsp;+\u0026thinsp;bacteria relative abundance:collection.\u003c/p\u003e \u003cp\u003eModels for shoots decomposition were as follows:\u003c/p\u003e \u003cp\u003eEq.\u0026nbsp;5\u0026thinsp;=\u0026thinsp;experimental variant+soil origin+collection+experimental variant:soil origin+experimental variant:collection+collection:soil origin\u003c/p\u003e \u003cp\u003eEq.\u0026nbsp;6\u0026thinsp;=\u0026thinsp;mass loss\u0026thinsp;~\u0026thinsp;fungal taxonomic richness\u0026thinsp;+\u0026thinsp;fungal taxonomic richness:experimental variant\u0026thinsp;+\u0026thinsp;fungal taxonomic richness:soil origin\u0026thinsp;+\u0026thinsp;fungal taxonomic richness:collection\u003c/p\u003e \u003cp\u003eEq.\u0026nbsp;7\u0026thinsp;=\u0026thinsp;mass loss\u0026thinsp;~\u0026thinsp;H-DR\u0026thinsp;+\u0026thinsp;H-DR:experimental variant\u0026thinsp;+\u0026thinsp;H-DR:soil origin\u0026thinsp;+\u0026thinsp;H-DR:collection\u003c/p\u003e \u003cp\u003eEq.\u0026nbsp;8\u0026thinsp;=\u0026thinsp;mass loss\u0026thinsp;~\u0026thinsp;bacteria relative abundance\u0026thinsp;+\u0026thinsp;bacteria relative abundance:experimental variant\u0026thinsp;+\u0026thinsp;bacteria relative abundance:soil origin\u0026thinsp;+\u0026thinsp;bacteria relative abundance:collection\u003c/p\u003e \u003cp\u003e.\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cp\u003eWe identified 25 different fungal morphotypes. The most common morphotypes were \u003cem\u003eFusarium\u003c/em\u003e sp., \u003cem\u003ePenicillium\u003c/em\u003e sp./\u003cem\u003eAspergillus\u003c/em\u003e sp., and \u003cem\u003eCladosporium\u003c/em\u003e sp., and the mean spores per sample was 31.08 with 40.38 SD. The mean number of bacterial cells per 100\u0026micro;m x 100\u0026micro;m square was 77.41 with 62.09 SD.\u003c/p\u003e \u003cp\u003eLeaves decomposition\u003c/p\u003e \u003cp\u003eAfter the first six weeks of the experiment, the mean mass loss was 34.67% (\u0026plusmn;\u0026thinsp;2.40% SD). After 12 weeks, it was 45.55% (\u0026plusmn;\u0026thinsp;3.41% SD), after 18 weeks, 49.78% (\u0026plusmn;\u0026thinsp;1.77% SD), and after 24 weeks, 52.31% (\u0026plusmn;\u0026thinsp;3.59% SD). Mean mass loss at the end of experiment was 52.82% (\u0026plusmn;\u0026thinsp;2.43% SD) in variant A (without filtering), 54.08% (\u0026plusmn;\u0026thinsp;3.86% SD) in variant B with filtering through a 10um filter, 52.04% (\u0026plusmn;\u0026thinsp;4.39% SD) in variant C with filtering through a 1.2um filter, and 53.64% (\u0026plusmn;\u0026thinsp;2.37% SD) in control (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e; Eq.\u0026nbsp;1).\u003c/p\u003e \u003cp\u003eOverall, fungal taxonomic diversity was responsible for a 10% mass loss increase (from 43% with one morphotype detected to 51% with 12 morphotypes). The addition of soil solution from invaded plots and control plots in interaction with fungal richness was responsible for 12% and 10% mass loss, respectively. After six weeks of the experiment, the fungal taxonomic diversity was responsible for a 14% decrease in the mass loss, and after 24 weeks for a 16% increase. In variant A, fungal diversity was responsible for 7% mass loss decrease, in variant B for a decrease, in variant C for 7% decrease, and in variant D for 2% increase (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e; Eq.\u0026nbsp;2).\u003c/p\u003e \u003cp\u003eWhen the abundance of DSM was higher than the abundance of hyaline mycelium decomposition ratio was also higher. Mass loss was 56% when the H-DR (hyaline minus dark ratio) was 10, and when the H-DR was 18, the mass loss ratio was 30%. There was no difference in the influence of mass loss between different soil origins in the case of the H-DR. After six weeks, the higher abundance of hyaline mycelium was responsible for a 21% decrease in the mass loss, and after 24 weeks, for a 13% decrease. In variant A, the higher abundance of hyaline mycelium was responsible for a 26% mass loss decrease, in variant B and D for a 27% mass loss decrease, and in variant C for an 18% mass loss decrease (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e; Eq.\u0026nbsp;3). Overall, bacterial relative abundance was responsible for 3% mass loss increase (from 48% with 20% of bacteria to 50% with 100% relative bacterial abundance). Bacterial relative abundance effects on the mass loss varied from negligible to weak (2% \u0026minus;\u0026thinsp;8%) in the interaction with other studied parameters. (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e;Eq.\u0026nbsp;4).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eShoots decomposition\u003c/p\u003e \u003cp\u003eDecomposition of shoots was slower than that of leaves, even with longer periods between collections. After the first 12 weeks of the experiment, the mean mass loss was 15.41% (\u0026plusmn;\u0026thinsp;2.36% SD), after 24 weeks, 18.45% (\u0026plusmn;\u0026thinsp;5.74% SD), after 36 weeks, 28.67% (\u0026plusmn;\u0026thinsp;7.39% SD), and after 48 weeks 30.66% (\u0026plusmn;\u0026thinsp;7.87% SD). Mean mass loss at the end of an experiment was 30.66% (\u0026plusmn;\u0026thinsp;7.87% SD) in variant A (without filtering), 54.08% (\u0026plusmn;\u0026thinsp;3.86% SD) in variant B with filtering through a 10um filter, 52.04% (\u0026plusmn;\u0026thinsp;4.39% SD) in variant C with filtering through 1.2um filter, and 53.64% (\u0026plusmn;\u0026thinsp;2.37% SD) in control (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e; Eq.\u0026nbsp;5).\u003c/p\u003e \u003cp\u003eOverall, fungal taxonomic richness was responsible for a 7% mass loss decrease (from 21% with one morphotype detected to 15% with 10 morphotypes). The addition of soil solution from invaded plots and control plots in interaction with fungal richness was responsible for a 6% and 5% mass loss decrease, respectively. After 12 weeks of the experiment, fungal taxonomic richness was responsible for a 14% decrease in the mass loss, and after 48 weeks for 18% increase. In variant A and C, fungal richness was responsible for 7% mass loss decrease, in variant B for a 10% decrease, and in variant D for 2% decrease (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e;Eq.\u0026nbsp;6).\u003c/p\u003e \u003cp\u003eWhen the abundance of dark septated mycelium (DSM) was higher than the abundance of hyaline mycelium, the decomposition ratio was also higher. Mass loss was\u003c/p\u003e \u003cp\u003e29% when the H-DR (hyaline minus dark ratio) was \u0026minus;\u0026thinsp;5, and when the H-DR was 20, the mass loss ratio was 9%. Higher abundance of hyaline mycelium was responsible for a 25% decrease in the decomposition in variants with invasive soil added, and a 20% decrease in variants with native soil added. After 12 weeks, the higher abundance of hyaline mycelium was responsible for a 21% decrease in the mass loss, and after 48 weeks for a 13% decrease. In variant A, the higher abundance of hyaline mycelium was responsible for a 21% mass loss decrease, in variant B for a 10% mass loss decrease, and in variants C and D for a 20% mass loss decrease (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e; Eq.\u0026nbsp;2).\u003c/p\u003e \u003cp\u003eOverall, bacterial relative abundance was responsible for a 12% mass loss decrease (from 22% with 20% of bacteria to 12% with 100% relative bacterial abundance). Soil origin had a negligible effect on the influence of the bacterial abundance on the mass loss. After 12 weeks of the experiment, bacterial relative abundance was responsible for a 13% decrease in the mass loss, and after 48 weeks for 7% increase. In all experimental variants, the abundance of bacteria had a similar negative effect (10% to 14% mass loss decrease, Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e; Eq.\u0026nbsp;5).\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eMass loss within different experimental variants\u003c/p\u003e \u003cp\u003eIn the current study, the mass loss grew faster than in earlier studies on \u003cem\u003eS. canadensis\u003c/em\u003e litter decay conducted in the terrestrial environments (e.g., Zhang et al., \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Nevertheless, the mass loss we obtained was close to the \u003cem\u003eS. canadensis\u003c/em\u003e shoot and leaf decomposition ratios reported by Anda et al. (\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) in the freshwater. This similarity was expected, as both our experiment and that of Anda \u003cem\u003eet al.\u003c/em\u003e eliminated the negative effects of drought on microbial activity through their experimental setups. On the other hand, we expected more factors in our study to affect the decomposition. First of all, our expectations regarding the impact of soil origin (invaded vs. control plots) were not confirmed. We observed only negligible differences between filtration variants of soil solution and variants of suspended soil (from \u003cem\u003eSolidago\u003c/em\u003e stands, and mesic meadows without \u003cem\u003eSolidago\u003c/em\u003e). Moreover, our data do not show any consistent trend in these relationships. For instance, after 36 weeks, variants with soil from non-invaded plots had slightly higher mass loss than those with soil solution from invaded ones, but after another 12 weeks, their effects were relatively similar. As shown by Kisło et al. (\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), the increasing cover of \u003cem\u003eS. canadensis\u003c/em\u003e on experimental plots does not affect the rate of leaf litter decomposition. Moreover, the main part of this study - different fractions of soil solutions expressed as soil solutions\u0026rsquo; filtering variants - turned out to be ineffective predictors of \u003cem\u003eS. canadensis\u003c/em\u003e litter decomposition as soil solution origin (i.e., invaded and non-invaded meadows).\u003c/p\u003e \u003cp\u003eAs regards soil origin, we observed slight differences between the decomposition speed within different filtration variants, but they are hard to interpret ecologically. They rather derive from interspecies litter quality (e.g., organs\u0026rsquo; thickness, or lignin concentration), which may exert a pronounced impact on the decomposition (Couˆteaux \u003cem\u003eet al.\u003c/em\u003e, 1995), than from ecological processes concerning the impact of saprotrophic biota. Moreover, this indicates a high degree of functional redundancy among \u003cem\u003eS. canadensis\u003c/em\u003e decomposers. More specifically, when one functional group was removed or underperformed, others were able to compensate and maintain decomposition processes. For instance, the elimination of soil-borne fungi allowed bacteria to occupy more environmental niches, thus degrading more types of substrates, since the difference in mass loss between variants A, B, and C was negligible. Moreover, in our samples, we observed \u003cem\u003eHarzia\u003c/em\u003e sp., a mycopathogenic fungus (Nguyen et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), which may have regulated the fungal population, and by that, the fungal impact on the decomposition, when other fungivorous organisms were missing.\u003c/p\u003e \u003cp\u003eVariant D (control), consisting solely of \u003cem\u003eS. canadensis\u003c/em\u003e leaves/shoots and their native biota, showed a mass loss level comparable to the other variants. This allows for a cautious hypothesis that the microbiota naturally inhabiting \u003cem\u003eS. canadensis\u003c/em\u003e tissues may contribute to the decomposition of its own litter. Endophyte-driven priming has already been demonstrated (Guerreiro \u003cem\u003eet al.\u003c/em\u003e, 2017), and our results may indicate a similar but not yet fully confirmed mechanism. Placed in the hologenome context (Moran and Sloan, \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2015\u003c/span\u003e), this would mean that the plant and its tissue-associated microbes form a unit capable of recycling nutrients embedded in the biomass. This link is particularly relevant for invasive species such as \u003cem\u003eS. canadensis\u003c/em\u003e, which have a high nutrient demand due to their reproduction strategy based on excessive diaspore production (Hua \u003cem\u003eet al.\u003c/em\u003e, 2007; Ye \u003cem\u003eet al.\u003c/em\u003e, 2018). Moreover, \u003cem\u003eS. canadensis\u003c/em\u003e represents a \u0026ldquo;try harder\u0026rdquo; strategy, characterized by an acquisitive pattern of resource use aimed at maximizing growth and reproductive output (Tecco et al., \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Because these nutrients are strongly accumulated in the leaves, conditions are favourable for the development of saprotrophs within the plant\u0026rsquo;s own tissues. This may explain both the similarity between our results and those of Anda et al. (\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) and the limited effect of soil origin or filtering. \u003cem\u003eS. canadensis\u003c/em\u003e may already carry saprotrophs capable of initiating decomposition, supporting nutrient return to the plant or its offspring.\u003c/p\u003e \u003cp\u003eBacteria and fungi have similar effects on the decomposition\u003c/p\u003e \u003cp\u003eUnexpectedly, the relative abundance of bacteria and fungal diversity showed a negative correlation with mass loss at the beginning of the experiment. Plant litter with \u003cem\u003eS. canadensis\u003c/em\u003e origin may be hard to degrade due to high lignin concentration in shoots (Wiatrowska et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Probably cuticle, as the natural barrier between inner plant tissues and the environment, needed the initial degradation process, as shown in (Logan \u003cem\u003eet al.\u003c/em\u003e, 2022). Moreover, \u003cem\u003eS. canadensis\u003c/em\u003e is known for producing a high diversity of secondary metabolites with different stability and potential impact on the decomposition (Zaimenko et al., \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Nevertheless, a high abundance of microorganisms that are not rapid decomposers may contribute to the residual mass, thereby slowing the overall rate of mass loss.\u003c/p\u003e \u003cp\u003eThe production of secondary metabolites in the cuticule-related tissues in \u003cem\u003eS. canadensis\u003c/em\u003e organs may be the reason for decreasing the impact of fungi and bacteria on the mass loss at the beginning of the experiment in both shoots and leaves. Notable inhibition by these groups\u0026rsquo; representatives was visible after six weeks of experiment in the case of leaf litter and after 12 and 24 weeks in the case of shoots, and we found a probable explanation for this phenomenon. Secondary metabolites of \u003cem\u003eS. canadensis\u003c/em\u003e may differ between shoots and leaves, or among different organs. Anžlovar et al. (\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) showed this experimentally: extracts from different organs had varying effects on \u003cem\u003eBotrytis cinerea\u003c/em\u003e growth. Studies by Reyes-\u0026Aacute;vila et al. (\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) and Jenner et al. (\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) have shown that two of the secondary metabolites present in \u003cem\u003eS. canadensis\u003c/em\u003e tissues, germacene D and limonene, have different decomposition ratios. Limonene decomposes fast, in up to seven days in the soil, and up to 40 days in the distilled water (Reyes-\u0026Aacute;vila et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), while more than 75% of germacene D survives 63 days of biodegradation (Jenner et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Moreover, we can expect that the substrate itself remains largely unavailable to saprotrophs, yet the aquatic environment becomes sufficiently enriched with dissolved organic compounds to stimulate bacterial cell division. Thus, bacterial abundance may increase even though the substrate is not yet undergoing intensive decomposition.\u003c/p\u003e \u003cp\u003eSince the current study focuses mainly on the fungal idiophase (spores), and sporulation sometimes requires specific conditions (Morton, \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1961\u003c/span\u003e), we also checked the impact of the trophophase (vegetative mycelium) on the decomposition. Despite the knowledge that fungi with a trophophorm of hyaline mycelium have fast metabolism and are effective degraders (i.e., Basidiomycetes (Pleurotus), Mucoromycetes; Cohen \u003cem\u003eet al.\u003c/em\u003e, 2002; Satari and Karimi, 2017), our results suggest the opposite dependence. Some plant-associated fungi with melanised hyphae, such as \u003cem\u003eCladosporium\u003c/em\u003e spp., show broad metabolic and functional diversity (Bensch et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Nguyen et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). However, this factor alone does not explain why our findings differ from those of the field study by Kisło et al. (\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), in which DSM had a clearly negative effect on mass loss. One possible explanation is that under constant high humidity and optimal temperature, DSM fungi may allocate more resources to faster trophic metabolism. Another possibility is that environmental conditions select for different taxonomic groups of DSM-associated decomposers in different settings. Unfortunately, our experiment could not determine which of these two possibilities is closer to the truth.\u003c/p\u003e \u003cp\u003eTaken together, these patterns indicate that bacteria and fungi exert broadly similar effects on the decomposition of both leaves and shoots, regardless of soil solution origin. This consistency suggests that external microbial communities play only a minor role in shaping early and mid-term decomposition dynamics in \u003cem\u003eS. canadensis\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eStudy limitations\u003c/p\u003e \u003cp\u003eOur experiment was conducted at room temperature, which removed the effect of varying environmental conditions but may have limited the activity of fungi requiring stratification for germination (Juge \u003cem\u003eet al.\u003c/em\u003e, 2002). We also detected fungal spores and nematodes in soil-filtration variants. Because their presence on leaves and shoots was more likely than filtration errors, we treated them as components of the endophytic/epiphytic biota of \u003cem\u003eS. canadensis\u003c/em\u003e. Only direct microscopy was used, as culturing would require partitioning the material and could introduce decomposition-rate biases. For the same reason, we excluded the metabarcoding approach. Using soil for experimental preparation also increased the risk of detecting dormant spores or eggs unrelated to \u003cem\u003eS. canadensis\u003c/em\u003e decomposition, potentially producing false metabarcoding signals. Moreover, the magnification used allows for a relative comparison of bacterial abundance in the sample, but without specific staining, some fraction of the bacteria may be underestimated. Moreover, some bacteria occur in colonies, and such colonies are likely to be analysed as single units.\u003c/p\u003e \u003cp\u003eFuture experiments should therefore include two storage variants-room temperature and conditions matching the litter collection site-to avoid limiting fungal stratification. Metabarcoding should be performed before the experiment (on fresh and dried litter) to confirm the presence or absence of non-endophytic biota. Additionally, using litter from plants grown under sterile conditions would help eliminate biases introduced by endophytic communities.\u003c/p\u003e"},{"header":"Conclusions","content":"\u003cp\u003e\u003cem\u003eSolidago canadensis\u003c/em\u003e is known for acting as an environmental engineer and changing the properties of the environment in the introduced range. When we add novel interactions with endophytic biota acting as efficient decomposers in the introduced range, as stated in Kisło et al. (\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2025\u003c/span\u003e) and in the current study, we obtain a view of a self-sufficient invader, and by that, fully adapted to modifying new areas. Our concern shall be especially important for lawmakers preparing local lists of invasive species with the highest impact on the environment and the lists of non-native species that should be eradicated from their secondary range. The second, more optimistic view of our results provides a valuable insight into the applied usage of \u003cem\u003eS. canadensis\u003c/em\u003e. Since its tissues contain biota that are sufficient decomposers, the effort put into the composting of this plant seems to be relatively low, thus with a relatively low cost. In this case, our results may be part of the opportunity to fight the invasion of this plant, since mowing its shoots twice a year, according to the instructions (Gala-Czekaj et al., \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), may be profitable for both farmers and companies selling bioproducts.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003eAcknowledgments\u003c/p\u003e\n\u003cp\u003eAuthor Contributions\u003c/p\u003e\n\u003cp\u003eAll authors contributed to the conception and design of the study. Kamil Kisło, Patryk Czortek, Anna Wiewiorowska, and Marta Wrzosek performed material preparation, data collection, and analysis. The first draft of the manuscript was written by Kamil Kisło, and all authors commented on previous versions. All authors read and approved the final manuscript.\u003c/p\u003e\n\u003cp\u003eData Availability\u003c/p\u003e\n\u003cp\u003eThe datasets generated and/or analyzed during the current study are available from the corresponding author upon reasonable request.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAkatov Vv, Akatova T v., Chefranov SG (2021) Impact of \u003cem\u003eSolidago canadensis\u003c/em\u003e L. on Species Diversity of Plant Communities at Different Spatial Scale. 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Pedosphere 26:567\u0026ndash;576. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/S1002-0160(15)60066-2\u003c/span\u003e\u003cspan address=\"10.1016/S1002-0160(15)60066-2\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"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":"plant-and-soil","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"plso","sideBox":"Learn more about [Plant and Soil](https://www.springer.com/journal/11104)","snPcode":"11104","submissionUrl":"https://submission.nature.com/new-submission/11104/3","title":"Plant and Soil","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Solidago canadensis, invasive plant, environmental engineer, fungi, bacteria, decomposition","lastPublishedDoi":"10.21203/rs.3.rs-8917351/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8917351/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eBackground and Aims\u003c/p\u003e \u003cp\u003e \u003cem\u003eSolidago canadensis\u003c/em\u003e is an invasive herb widespread across Eurasia and Australia, known to decompose faster than native species. However, the mechanisms driving its decomposition, particularly the role of associated microorganisms, remain poorly understood. This study aimed to assess the influence of specific components of soil biota (bacteria and fungi) from invaded and non-invaded mesic meadows on the decomposition of \u003cem\u003eS. canadensis\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eMethods\u003c/p\u003e \u003cp\u003eWe conducted an \u003cem\u003eex situ\u003c/em\u003e decomposition experiment using shoots and leaves of \u003cem\u003eS. canadensis\u003c/em\u003e. Soil solutions from invaded and non-invaded meadows were prepared and passed through syringe filters of varying pore sizes (variants A\u0026ndash;D). We assessed primary decomposer diversity under a light microscope and analysed mass loss using generalized linear mixed models.\u003c/p\u003e \u003cp\u003eResults\u003c/p\u003e \u003cp\u003eDifferences among filtration treatments were minimal, and soil solution origin (invaded vs. non-invaded meadows) had negligible effects on mass loss. Leaves and shoots decomposed at similar rates across all variants. These outcomes indicate that external microbial inputs played no detectable role in driving decomposition.\u003c/p\u003e \u003cp\u003eConclusion\u003c/p\u003e \u003cp\u003eOur findings suggest that \u003cem\u003eS. canadensis\u003c/em\u003e operates as a largely self-contained holobiont, harbouring an internal consortium of primary decomposers capable of sustaining decomposition independently of external soil communities. This highlights the functional redundancy of surrounding soil biota and may contribute to enhanced nutrient cycling in invaded ecosystems. Such a strategy aligns with invasion facilitation frameworks, including the novel weapon hypothesis, illustrating how \u003cem\u003eS. canadensis\u003c/em\u003e can exploit new environments. Additionally, its consistently efficient \u003cem\u003eex situ\u003c/em\u003e decomposition suggests strong potential for valorizing harvested biomass through composting or biogas production.\u003c/p\u003e","manuscriptTitle":"Solidago canadensis and its endophytes as a self-degrading system during ex situ decomposition","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-03-02 16:03:25","doi":"10.21203/rs.3.rs-8917351/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Major revisions","date":"2026-05-04T04:42:21+00:00","index":"","fulltext":""},{"type":"reviewerAgreed","content":"","date":"2026-02-27T00:29:43+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2026-02-26T07:43:16+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"Plant and Soil","date":"2026-02-20T05:51:51+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2026-02-20T01:52:23+00:00","index":"","fulltext":""},{"type":"submitted","content":"Plant and Soil","date":"2026-02-19T06:54:38+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"plant-and-soil","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"plso","sideBox":"Learn more about [Plant and Soil](https://www.springer.com/journal/11104)","snPcode":"11104","submissionUrl":"https://submission.nature.com/new-submission/11104/3","title":"Plant and Soil","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"dc297dfc-d9ff-4cd3-ac65-ada77dcecad2","owner":[],"postedDate":"March 2nd, 2026","published":true,"recentEditorialEvents":[{"type":"decision","content":"Major revisions","date":"2026-05-04T04:42:21+00:00","index":"","fulltext":""}],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-05-18T14:41:08+00:00","versionOfRecord":[],"versionCreatedAt":"2026-03-02 16:03:25","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8917351","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8917351","identity":"rs-8917351","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}