Divergent responses of rhizosphere soil phosphorus fractions and biological features of Salix psammophila to fertilization strategies under cadmium contamination.

Soil oligotrophy in areas heavily contaminated with heavy metals poses a significant challenge to vegetation establishment and phytoremediation processes. Phosphorus (P) cycling plays a critical role in global biogeochemical cycles, but there is limited understanding of its response to varying fertilization strategies and its correlation with phytoremediation effectiveness. This study primarily investigated the effects of various fertilization strategies, including nitrogen (N, 300 mg·kg-1), P (100 mg·kg-1), NP (combined N and P at 300 mg·kg-1 and 100 mg·kg-1, respectively), and HP (high P, 300 mg·kg-1) application, on rhizosphere soil P fractions and P-solubilizing microbial community (harboring phoD and phoC genes, respectively) of Salix psammophila under cadmium contamination. Application of NP significantly enhanced plant growth and cadmium accumulation, whereas HP inhibited cadmium bioaccumulation but promoted its translocation. Compared to untreated soil, N application promoted P cycling, leading to increases of 141.9 %, 60.4 %, and 10.3 % in Resin-Pi, diluted HCl-Pi, and conc.HCl-Pi, respectively. P application decreased organic phosphorus (Po) fractions by 24.4 % - 225.8 %, but N incorporation mitigated the declining trend in Po and augmented alkaline phosphatase activity. Fertilization strategies significantly regulated phoC- or phoD-harboring bacterial community structure, but their differential nutrient demands resulted in distinct responses. The phoD-harboring bacteria exhibited higher diversity and network complexity, with numerous biomarkers and fertilizer-sensitive OTUs discovered across treatments. Structural equation modeling (SEM) analysis indicated that phytoremediation efficiency was directly affected by Pi fractions, and phoD-harboring bacteria exhibited stronger associations with Pi fractions than phoC-harboring bacteria. In conclusion, our results reveal potential pathways through which fertilization strategies influence phytoremediation by affecting the structure of P-solubilizing microbial community. Furthermore, our study emphasizes the importance of combined N and P application in promoting Cd accumulation in plants, with high P levels appearing as an ideal fertilization strategy for phytoremediation targeting the harvest of aboveground biomass.

Enriched rhizospheric functional microbiome may enhance adaptability of Artemisia lavandulaefolia and Betula luminifera in antimony mining areas.

Dominant native plants are crucial for vegetation reconstruction and ecological restoration of mining areas, though their adaptation mechanisms in stressful environments are unclear. This study focuses on the interactions between dominant indigenous species in antimony (Sb) mining area, Artemisia lavandulaefolia and Betula luminifera, and the microbes in their rhizosphere. The rhizosphere microbial diversity and potential functions of both plants were analyzed through the utilization of 16S, ITS sequencing, and metabarcoding analysis. The results revealed that soil environmental factors, rather than plant species, had a more significant impact on the composition of the rhizosphere microbial community. Soil pH and moisture significantly affected microbial biomarkers and keystone species. Actinobacteria, Proteobacteria and Acidobacteriota, exhibited high resistance to Sb and As, and played a crucial role in the cycling of carbon, nitrogen (N), phosphorus (P), and sulfur (S). The genes participating in N, P, and S cycling exhibited metabolic coupling with those genes associated with Sb and As resistance, which might have enhanced the rhizosphere microbes' capacity to endure environmental stressors. The enrichment of these rhizosphere functional microbes is the combined result of dispersal limitations and deterministic assembly processes. Notably, the genes related to quorum sensing, the type III secretion system, and chemotaxis systems were significantly enriched in the rhizosphere of plants, especially in B. luminifera, in the mining area. The phylogenetic tree derived from the evolutionary relationships among rhizosphere microbial and chloroplast whole-genome resequencing results, infers both species especially B. luminifera, may have undergone co-evolution with rhizosphere microorganisms in mining areas. These findings offer valuable insights into the dominant native rhizosphere microorganisms that facilitate plant adaptation to environmental stress in mining areas, thereby shedding light on potential strategies for ecological restoration in such environments.

Modulating phyllosphere microbiome structure and function in Loropetalum chinense and Osmanthus fragrans: The impact of foliar dust and heavy metals.

Trees can effectively capture airborne particles and improve air quality. However, the specific response of phyllosphere microbiome (PMo) in different plant species to particulate matter (PM) and the heavy metals it contains are not yet fully understood. In this study, we investigated the impact of PM on the diversity and function of PMo in Loropetalum chinense and Osmanthus fragrans trees grown in industrial and clean zones with varying levels of PM pollution. Our findings revealed that leaf dust had a significant negative effect on microbial richness, with O. fragrans exhibiting higher microbial diversity than L. chinense. The dominant phylum of phyllosphere bacteria in all samples was Proteobacteria, and the dominant genera were Stenotrophomonas and Delftia. The relative abundance of these genera varied significantly among plant species and regions. Our results showed that PM had a significant impact on the community composition of PMo, with the presence of heavy metals exerting a greater effect than particle size. Moreover, the foliar microbial community of plants grown in industrial zones exhibited significantly higher metabolic functions related to stress resistance and disease resistance compared to plants in control zones. These findings highlight the structural and functional responses of PMo to PM and indicate their potential for enhancing plant adaptation to environmental stress.

Evaluating the role of rhizosphere microbial home-field advantage in Betula luminifera adaptation to antimony mining areas.

It has been established that the coevolution of plants and the rhizosphere microbiome in response to abiotic stress can result in the recruitment of specific functional microbiomes. However, the potential of inoculated rhizosphere microbiomes to enhance plant fitness and the inheritance of adaptive traits in subsequent generations remains unclear. In this study, cross-inoculation trials were conducted using seeds, rhizosphere microbiome, and in situ soil collected from areas of Betula luminifera grown in both antimony mining and control sites. Compared to the control site, plants originating from mining areas exhibited stronger adaptive traits, specifically manifested as significant increases in hundred-seed weight, specific surface area, and germination rate, as well as markedly enhanced seedling survival rate and biomass. Inoculation with mining microbiomes could enhance the fitness of plants in mining sites through a "home-field advantage" while also improving the fitness of plants originating from control sites. During the initial phase of seedling development, bacteria play a crucial role in promoting growth, primarily due to their mechanisms of metal resistance and nutrient cycling. This study provided evidence that the outcomes of long-term coevolution between plants and the rhizosphere microbiome in mining areas can be passed on to future generations. Moreover, it has been demonstrated that transgenerational inheritance and rhizosphere microbiome inoculation are important factors in improving the adaptability of plants in mining areas. The findings have important implications for vegetation restoration and ecological environment improvement in mining areas.

Facilitated remediation of heavy metals contaminated land using Quercus spp. with different strategies: Variations in amendments and experiment periods.

Phytoremediation using trees combined with soil amendments has gained much attention for its highly cost-effective trait. In natural field conditions, however, the results may not reflect the true performance of amendments based on short-term laboratory studies. In this three-year field trial, various soil amendments such as rice straw biochar, palygorskite, a combined biochar of rice straw biochar and palygorskite, and hydroxyapatite were used to systematically study the potential of the low-accumulator (Quercus fabri Hance) and high-accumulator (Quercus texana Buckley) for cadmium (Cd) and zinc (Zn) to remediate severely contaminated soils. Soil amendments enhanced the dendroremediation capacity of Quercus as the growth period prolonged. In 2021, the rice straw biochar treatment increased Cd and Zn accumulation by 1.76 and 2.09 times in Q. fabri, respectively, compared to the control. Cd and Zn accumulation increased to 1.78 and 2.10 times, respectively, under combined biochar treatment for Q. texana compared to the control. Metals accumulation was mainly enhanced by soil amendments through increasing the growth biomass of Q. fabri and improving the biomass and bioconcentration ability of Q. texana. Overall, soil amendments effectively improved the phytoremediation efficiency of Quercus in the long term, and selecting suitable amendments should be fully considered in phytoremediation.

Microbial communities in tree root-compartment niches under Cd and Zn pollution: Structure, assembly process and co-occurrence relationship.

Woody plants have showed great potential in remediating severely contaminated soils by heavy metals (HMs) due to their cost-efficient and ecologically friendly trait. It is believed the root-associated microbiota plays a vital role in phytoremediation for HMs. However, the ecological process controlling the assembly and composition of tree root-associated microbial communities under HMs stress remains poorly understood. Herein, we profiled the bulk soil, rhizosphere and endosphere microbial communities of trees growing in heavily Cd and Zn polluted soil. The microbiota was gradually filtered from bulk soil to the tree roots and was selectively enriched in roots with specific taxa, such as Proteobacteria and Ascomycota. The microbial community assembly along the soil-root continuum was mainly controlled by deterministic processes from bulk soil to the endosphere, with the normalized stochasticity ratio (NST) indices of 67.16-31.05 % and 30.37-15.02 % for bacteria and fungi, respectively. Plant selection pressure sequentially increased from bulk soil to rhizosphere to endosphere, with the reduced bacterial alpha diversity accompanying the consequently reduced complexity of the co-occurrence network. Together, the findings provide new evidence for horizontal transmission of endophytic microbiome from soil to the host, which can shed light on the future screening and application of microbial-assisted phytoremediation.

Effects of on- and off-year management practices on the soil organic C fractions and microbial community in a Moso bamboo (Phyllostachys edulis) forest in subtropical China.

On- and off-year management practices are usually adopted in Moso bamboo (Phyllostachys edulis) forests to achieve higher productivity. However, little is known about the effects of these management practices on soil C sequestration and microbial community structure. In the present study, soil nutrient content, organic C fractions, and bacterial and fungal communities were comparatively investigated in on- and off-year bamboo stands. The results showed that soil organic C (SOC), alkali-hydrolyzable N (AN), and available P (AP) in the on-year were significantly lower (p ≤ 0.05) than those in the off-year. Among the different soil organic C fractions, easily oxidizable organic C (EOC), microbial biomass C (MBC), Ca-bound SOC (Ca-SOC), and Fe/Al-bound SOC (Fe/Al-SOC) also had significantly higher contents in the off-year than in the on-year, with MBC and EOC decreasing by 56.3% and 24.5%, respectively, indicating that both active and passive soil organic C pools increased in the off-year. However, the alpha diversities of both soil bacteria and fungi were significantly lower in the off-year soils than in the on-year soils. The bacterial taxa Actinobacteria, Planctomycetes, WPS-2, Acidothermus, Candidatus_Solibacter, Burkholderia-Caballeronia-Paraburkholderia, and Candidatus_Xiphinematobacter were increased in off-year soils relative to on-year soils. Meanwhile, fungal taxa Ascomycota, Mortierella, Hypocrea, Cryptococcus, Clitopilus, and Ceratocystis were significantly increased in on-year soils. Soil pH, SOC, AP, MBC, EOC, and Ca-SOC were significantly correlated with bacterial and fungal communities, with soil pH being the most important driving factor for the shift in bacterial and fungal communities. Our findings showed that the studied bamboo forest possessed an inherent restorative ability in the off-year, which can reverse the soil nutrient and C depletion in the on-years and ensure soil fertility in the long term.

Corrigendum: Effects of on- and off-year management practices on the soil organic C fractions and microbial community in a Moso bamboo (Phyllostachys edulis) forest in subtropical China.

[This corrects the article DOI: 10.3389/fpls.2022.1020344.].

Intercropping improves heavy metal phytoremediation efficiency through changing properties of rhizosphere soil in bamboo plantation.

Moso bamboo is considered a potential species for heavy metal (HM) phytoremediation; however, the effect of intercropping on rhizosphere and phytoextraction remains to be elucidated. We comparatively investigated rhizobacteria, soil properties, and phytoextraction efficiency of monoculture and intercropping of Moso bamboo and Sedum plumbizincicola in Cu/Zn/Cd-contaminated soil. Compared with monocultures, intercropping increased the bacterial α-diversity indices (Shannon, Chao1) and the number of biomarkers. Intercropping reduced the contents of soil organic matter (SOM), available nutrients, and Cd and Cu in rhizosphere soils, and reduced the Cd and Zn contents in tissues of sedum. By contrast, Cd and Zn contents in tissues of bamboo increased, and the increase of organic acid in root exudates from intercropping could facilitate the HM absorption. The total amount of Cu, Zn, and Cd removed from the soil in intercropping system was 1.2, 1.9, and 1.8 times than those in monoculture bamboo, respectively. The abundances of Proteobacteria, Acidobacteria, Verrucomicrobia and Actinobacteria were higher in intercropping, playing an important role in soil nutrient cycles and HM remediation. These bacterial communities were closely correlated (P < 0.01) with SOM, available nitrogen, available phosphorus, and HMs. The results suggested this intercropping pattern can increase HM removal efficiency from polluted soils.

Understanding variations in soil properties and microbial communities in bamboo plantation soils along a chromium pollution gradient.

With high biomass productivity and resistance to heavy metals (HM) stress, bamboo has strong potential for HM phytoremediation. However, few studies have been conducted under field conditions to explore changes in soil physicochemical and microbial properties of bamboo forests with HM-contaminated soils. This study established bamboo (Phyllostachys praecox) plantations in five Cr-contaminated sites with different pollution levels (low, L; low-moderate, LM; moderate, M; moderate-high, MH; and high, H). We determined soil chemical properties, total and available Cr content, as well as bacterial and fungal community structures from 0 to 20 cm depth along the pollution gradient, and evaluated their interactions. The results revealed a corresponding decrease in soil pH, alkali-hydrolysable N (AN), along with urease and sucrase activities, as Cr pollution increased. In contrast, total organic carbon (TOC) increased with increasing Cr pollution. Soil available P (AP) and acid phosphatase activity did not differ significantly. Different pollution level resulted in distinct bacterial and fungal communities, with Proteobacteria, Acidobacteria, Actinobacteria, Ascomycota, and Basidiomycota being the dominant phyla across the five bamboo soils. Both total Cr (TCr) and HCl-extractable Cr (ACr) negatively correlated with alpha indices (Chao1 and Shannon) for bacteria but not for fungi, indicating that the latter is more resistant to Cr pollution. Decrease in soil pH and increase in TCr and ACr from L to H were closely related to the shift of bacterial and fungal communities. These changes reduced soil N and C cycles. Our findings suggest that improving soil acidic conditions and N availability enhances carbon and nitrogen cycles via altering soil microbial structure and activities. This, in turn, can increase phytoremediation efficiency in the bamboo ecosystem.

Soil carbon and associated bacterial community shifts driven by fine root traits along a chronosequence of Moso bamboo (Phyllostachys edulis) plantations in subtropical China.

Moso bamboo (Phyllostachys edulis) is widely considered to be effective in capturing and sequestering atmospheric C, but the long-term effects of extensive management strategies on soil organic carbon (SOC), bacterial communities, fine root (FR, ø ≤ 2 mm) traits, and their inherent connection remain unclear. In this study, we simultaneously measured the SOC content of the bulk and rhizosphere soil fractions, the aggregate stability, the chemical composition of SOC (solid-state 13C nuclear magnetic resonance [NMR]), the bacterial community structure in the rhizosphere, and the FR morphological traits including biomass, specific root length (SRL), and root length density (RLD) along a chronosequence (stand age of 19, 37, and 64 years) of extensively managed Moso bamboo plantations and in an adjacent secondary forest as a control. The organic C content in both the rhizosphere and bulk soil increased rapidly with plantation age in the 0-20- and 20-40-cm soil layers, accompanied by an increase in the aggregate stability. FR traits including biomass, SRL, and RLD also increased continuously in response to soil C:N:P stoichiometry. All of these traits were significantly correlated with SOC, occluded particulate organic C (oPOC), and mineral-associated organic C (MOC), suggesting that FR traits could drive the soil C sequestration with the plantation age. Further analysis indicated that the microbial biomass C (MBC) content, MBC/total organic carbon (TOC) ratio, and bacterial abundance decreased with the plantation age, and the shift from soil oligotrophy to copiotrophy bacteria were mainly driven by changes in FR traits and SOC properties. Such a reassembly of bacterial communities combined with an increase in root biomass is favorable for the accumulation of stable C functional groups (alkyl C or aromatic C). Our findings indicate that extensive management regimes of Moso bamboo plantations could promote long-term soil C sequestration especially in the rhizosphere by promoting the formation of soil aggregates and organic-mineral complexes and by shifting bacterial community composition, and that these changes can be inferred through changes in the FR traits.

Deciphering the rhizosphere microbiome of a bamboo plant in response to different chromium contamination levels.

Bamboo has been considered a potential plant species for phytoremediation due to its high biomass and heavy metal (HM) resistance. However, little is known about the interactions between bamboo and soil microbial activities in HM-contaminated soils. Here, we investigated the characteristics of microbial communities in the rhizosphere soil of Lei bamboo (Phyllostachys praecox) along a chromium (Cr) gradient. We found that the soil Cr content was positively correlated with the total organic carbon (TOC) and HCl-extractable Cr but negatively correlated with the pH and bacterial and fungal Shannon indices. Proteobacteria and Ascomycota predominated in the bamboo rhizosphere under Cr pollution. A co-occurrence network showed that two of the most Cr-sensitive bacterial genera and keystone taxa were from the Acidobacteria, indicating that this phylum can be as an indicator for the studied Cr-polluted soils. Redundancy analysis revealed that both the soil bacterial and fungal community compositions were significantly correlated (p < 0.05) with Cr, pH, TOC, alkali-hydrolysable N (AN), and available phosphorus (AP). The increase in TOC as the Cr content increased can be ascribed to an adverse Cr effect on the soil microflora, probably because the microbial biomass was less effective in mineralizing soil C under Cr-polluted conditions.

Bamboo - An untapped plant resource for the phytoremediation of heavy metal contaminated soils.

Phytoremediation is a green technology used for the remediation of heavy metal soils. However, up to now, very few plants are known to be both hyperaccumulators and fast-growers. In contrast, some non-hyperaccumulators, which possess lower extraction capacities than hyperaccumulators, are fast-growing species with much higher total biomass yields and are potential alternative phytoremediators. Bamboo is a taxonomic group comprised of 1439 species that are mostly distributed in the tropics and subtropics. Although limited studies on bamboo for phytoremediation, recent studies have shown that some bamboo species have high ability to adapt to metalliferous environments and a high capacity to absorb heavy metals. Bamboo tissues in the rhizome and culm can accumulate a large amount of heavy metals that mainly accumulate in the cell wall, vacuole, and cytoplasm. Certain bamboo species such as moso bamboo, Phyllostachys praecox, have been shown to have a high endurance in metal contaminated soils, enabling a considerable uptake and accumulation of heavy metals. However, excessive concentrations of heavy metals may cause oxidative stress and damage bamboo plants. Therefore, several management strategies have been developed to improve the phytoremediation ability of bamboo species, including the selection of tolerant bamboo species, intercropping with hyperaccumulators, fertilization applications, and employment of chelate in soil. This review demonstrates that bamboo species, which have high biomass productivity, short rotation, and high economic value, can be used for phytoremediation. However, the mechanisms of heavy metal uptake, transport, sequestration, and detoxification of different bamboo species require urgent investigation.

Responses of soil nutrients and microbial communities to intercropping medicinal plants in moso bamboo plantations in subtropical China.

Bamboo forests are one of the most important forest resources in subtropical China. A pure, single-layer bamboo forest is considered an optimal habitat for intercropping medicinal herbs. Soil microorganisms have an important role in various ecological processes and respond quickly to environmental changes. However, changes in soil nutrients and microbial communities associated with agroforestry cultivation methods remain poorly documented. In the present study, a pure moso bamboo (Phyllostachys edulis) forest (Con) and three adjacent moso bamboo-based agroforestry (BAF) systems (moso bamboo-Paris polyphylla (BP), moso bamboo-Tetrastigma hemsleyanum (BT) and moso bamboo-Bletilla striata (BB)) were selected; and their soil chemical properties and bacterial communities were studied and compared to evaluate the effects of agroforestry on soil bacterial communities and the relationship between soil properties and bacterial communities in BAF systems. Results showed that compared with soils under the Con, soils under the BAF systems had more (p < 0.05) soil organic carbon (SOC) and available nitrogen (AN) but lower (p < 0.05) pH and available potassium (AK). In addition, compared with the Con system, the BB and BT systems had significantly greater (p < 0.05) available phosphorus (AP). Compared with that in the Con system, the Shannon index in the BAF systems was significantly greater (p < 0.05), but the Chao1 index not different. On the basis of relative abundance values, compared with the Con soils, the BAF soils had a significantly greater abundance of (p < 0.05) Bacteroidetes and Planctomyces but a significantly lower abundance of (p < 0.05) Verrucomicrobia, Gemmatimonadetes and Candidatus Xiphinematobacter. Moreover, compared with the Con system, the BB and BT systems had a greater (p < 0.05) abundance of Actinobacteria, Rhodoplanes, Candidatus Solibacter and Candidatus Koribacter. Redundancy analysis (RDA) revealed that soil pH, SOC and AP were significantly correlated with bacterial community composition. Results of this study suggest that intercropping medicinal herbs can result in soil acidification and potassium (K) depletion; thus, countermeasures such as applications of K fertilizer and alkaline soil amendments are necessary for BAF systems.

Changes of Root Endophytic Bacterial Community Along a Chronosequence of Intensively Managed Lei Bamboo (Phyllostachys praecox) Forests in Subtropical China.

Endophytic bacteria widely exist inside plant tissues and have an important role in plant growth and development and the alleviation of environmental stress. However, little is known about the response of root-associated bacterial endophytes of Lei bamboo (Phyllostachys praecox) to intensive management, which is a common management practice for high bamboo shoot production in subtropical China. In this study, we comparatively investigated the root endophytic bacterial community structures in a chronosequence of intensively managed (5a, 10a, 15a, and 20a) and extensively managed plantations (as control, Con). The results showed that endophytic Proteobacteria was the dominant bacterial phylum in the bamboo roots. Intensive management significantly increased (p < 0.05) the bacterial observed species and Chao1 (except 5a) indices associated with bamboo roots. The relative abundances of Firmicutes, Bacteroidetes, and Actinobacteria (except 15a) in the intensively managed bamboo roots significantly increased (p < 0.05) compared with those in Con, while the relative abundance of Proteobacteria significantly decreased in intensively managed bamboo roots (p < 0.05). The phyla Proteobacteria, Actinobacteria, Bacteroidetes, and Firmicutes were the biomarkers in Con, 5a, 15a, and 20a, respectively. Redundancy analysis (RDA) showed that soil alkali-hydrolysable N (AN), available phosphorus (AP), available K (AK), and total organic carbon (TOC) were significantly correlated (p < 0.05) with the bacterial community compositions. Our results suggest that the root endophytic microbiome of Lei bamboo was markedly influenced by intensive management practices, and the available nutrient status could be the main driving factor for such shifts. Although heavy fertilization in the intensive management system increased the diversity indices, the rapid changes in root endophyte communities and their relevant functions might indicate a high risk for sustainable management.