Arsenic Hyperaccumulator Pteris Vittata Research Articles

Enterobacter sp. E1 increased arsenic uptake in Pteris vittata by promoting plant growth and dissolving Fe-bound arsenic

Microbes affect arsenic accumulation in the arsenic-hyperaccumulator Pteris vittata, but the associated molecular mechanism remains uncertain. Here, we investigated the effect of Enterobacter sp. E1 on arsenic accumulation by P. vittata. Strain E1 presented capacities of arsenate [As(V)] and Fe(III) reduction during cultivation. In the pot experiment with P. vittata, the biomass, arsenic content, and chlorophyll content of P. vittata significantly increased by 30.03%, 74.9%, and 112.1%, respectively. Strikingly, the water-soluble plus exchangeable arsenic (WE-As) significantly increased by 52.05%, while Fe-bound arsenic (Fe–As) decreased by 29.64% in the potted soil treated with strain E1. The possible role of activation of arsenic by strain E1 was subsequently investigated by exposing As(V)-absorbed ferrihydrite to the bacterial culture. Speciation analyses of As showed that strain E1 significantly increased soluble levels of As and Fe and that more As(V) was reduced to arsenite. Additionally, increased microbial diversity and soil enzymatic activities in soils indicated that strain E1 posed few ecological risks. These results indicate that strain E1 effectively increased As accumulation in P. vittata mainly by promoting plant growth and dissolving soil arsenic. Our findings suggest that As(V) and Fe(III)-reducer E1 could be used to enhance the phytoremediation of P. vittata in arsenic-contaminated soils.

Intercropped Amygdalus persica and Pteris vittata applied with additives presents a safe utilization and remediation mode for arsenic-contaminated orchard soil

Intercropping the arsenic (As) hyperaccumulator Pteris vittata with fruit trees can safely yield peaches in As-polluted orchards in South China. However, the soil As remediation effects and the related mechanisms of P. vittata intercropped with peach trees with additives in the north temperate zone have rarely been reported. A field experiment was conducted to systematically study the intercropping of peach (Amygdalus persica) with P. vittata with three additives [calcium magnesium phosphate (CMP), ammonium dihydrogen phosphate (ADP), and Stevia rebaudiana Bertoni residue (SR)] in a typical As-contaminated peach orchard surrounding a historical gold mine in Pinggu County, Beijing City. The results showed that compared with monoculture (PM) and intercropping without addition (LP), the remediation efficiency of P. vittata intercropping was significantly increased by 100.9 % (CMP) to 293.5 % (ADP). CMP and ADP mainly compete with available As (A-As) adsorbed to the surface of Fe-Al oxide through PO43−, while SR might activate A-As by enhancing dissolved organic carbon (DOC) in P. vittata rhizospheres. The photosynthetic rates (Gs) of intercropped P. vittata were significantly positively correlated with pinna As. The intercropping mode applied with the three additives did not obviously affect fruit quality, and the net profit of the intercropping mode (ADP) reached 415,800 yuan·ha−1·a−1. The As content in peaches was lower than the national standard in the intercropping systems. Comprehensive analysis showed that A. persica intercropped with P. vittata applied with ADP is better than other treatments in improving risk reduction and agricultural sustainability. In this study, a theoretical and practical basis is provided for the safe utilization and remediation of As-contaminated orchard soil in the north temperate zone.

Removal of Methylene Blue from Wastewater by Waste Roots from the Arsenic-Hyperaccumulator Pteris vittata: Fixed Bed Adsorption Kinetics

Phytoremediation of arsenic-contaminated water was successfully conducted by means of the perennial fern Pteris vittate, which is an arsenic-hyperaccumulator plant able to grow in hydroponic cultures. In order to avoid the costs linked to the disposal of As-contaminated biomass, in this work, Pteris vittata waste roots were tested as a low-cost bio-adsorbent for the removal of methylene blue (MB) from water in a fixed-bed adsorption configuration. As a matter of fact, methylene blue can negatively impact the growth and health of algae and plants by blocking light from reaching them in water, which can alter their normal biological processes. Previous works have already shown the potentiality of such material toward the uptake of methylene blue; however, all the studies conducted were just focused on batch-mode experiments. In this work, column runs were carried out at 20 °C, evaluating the bed void fraction for each test and hence estimating the apparent density of the material (300 g/L). The breakthrough curves collected were fitted by means of a mathematical model based on the linear driving force (LDF) approximation to obtain information on the mass transfer mechanism occurring in the system. A relation for the product between the LDF mass transfer coefficient and the solid specific surface (kLDFas) with respect to the Reynolds (Re) dimensionless number was obtained (kLDFas=0.45Re). The range of validity of such expression was Re<0.025. Its applicability was deeply discussed: in such conditions, the technology is ready to be tested at larger scales.

Arsenic induced plant growth by increasing its nutrient uptake in As-hyperaccumulator Pteris vittata: Comparison of arsenate and arsenite

Arsenic-hyperaccumulator Pteris vittata is efficient in taking up arsenate (AsV) and arsenite (AsIII), however, their impacts on P. vittata growth and nutrient uptake remain unclear. The uptake of AsV and AsIII, their influences on nutrient uptake and plant biomass, and As speciation were investigated in P. vittata after exposing to 5 or 50 μM AsV or AsIII for 12 d under hydroponics. The results show that AsV uptake in P. vittata was 1.2 times more efficient than AsIII, corresponding to 1.7–2.1 fold greater biomass than the control at 50 μM As. While AsV was dominant in the roots at ∼60%, AsIII was more dominant in the fronds at ∼70% in all treatments. Macronutrients P, K, Ca, and S were increased by 118−185% at 50 μM As, with greater uptake of micronutrients Fe, Mn, Cu, and Zn at 5 μM As. Further, positive correlations between P. vittata biomass and its As contents (r = 0.97), and P. vittata biomass and its S, Mg, P, or Ca contents (r = 0.70–0.98) were observed. Our results suggest that its increased nutrient uptake probably enhanced P. vittata growth under As exposure.

Arsenic hyperaccumulator Pteris vittata shows reduced biomass in soils with high arsenic and low nutrient availability, leading to increased arsenic leaching from soil

Plant-soil interactions affect arsenic and nutrient availability in arsenic-contaminated soils, with implications for arsenic uptake and tolerance in plants, and leaching from soil. In 22-week column experiments, we grew the arsenic hyperaccumulating fern Pteris vittata in a coarse- and a medium-textured soil to determine the effects of phosphorus fertilization and mycorrhizal fungi inoculation on P. vittata arsenic uptake and arsenic leaching. We investigated soil arsenic speciation using synchrotron-based spectromicroscopy. Greater soil arsenic availability and lower nutrient content in the coarse-textured soil were associated with greater fern arsenic uptake, lower biomass (apparently a metabolic cost of tolerance), and arsenic leaching from soil, due to lower transpiration. P. vittata hyperaccumulated arsenic from coarse- but not medium-textured soil. Mass of plant-accumulated arsenic was 1.2 to 2.4 times greater, but aboveground biomass was 74% smaller, in ferns growing in coarse-textured soil. In the presence of ferns, mean arsenic loss by leaching was 195% greater from coarse- compared to the medium-textured soil, and lower across both soils compared to the absence of ferns. In the medium-textured soil arsenic concentrations in leachate were higher in the presence of ferns. Fern arsenic uptake was always greater than loss by leaching. Most arsenic (>66%) accumulated in P. vittata appeared of rhizosphere origin. In the medium-textured soil with more clay and higher nutrient content, successful iron scavenging increased arsenic release from soil for leaching, but transpiration curtailed leaching.

Geographical distribution of As-hyperaccumulator Pteris vittata in China: Environmental factors and climate changes

Understanding the distribution of hyperaccumulators helps to implement more efficient phytoremediation strategies of contaminated sites, however, limited information is available. Here, we investigated the geographical distribution of the first-known arsenic-hyperaccumulator Pteris vittata in China and the key factors under two climate change scenarios (SSP 1-2.6 and SSP 5-8.5) at two time points (2030 and 2070). Species distribution model (MaxEnt) was applied to examine P. vittata distribution based on 399 samples from field surveys and existing specimen records. Further, among 23 environmental factors, 11 variables were used in the MaxEnt model, including temperature, precipitation, elevation, soil property, and UV-B radiation. The results show that P. vittata can grow in ~23% of the regions in China. Specifically, it is mainly distributed in 11 provinces of southern China, including Hainan, Guangdong, Guangxi, Yunnan, Guizhou, Hunan, Hubei, Jiangxi, Fujian, Zhejiang, and Jiangsu. Besides, eastern Sichuan, and southern Henan, Shaanxi, and Anhui are suitable for P. vittata growth. Under two climate change scenarios, P. vittata distribution in China would decrease by ~5.76-7.46 × 104 km2 in 2030 and ~3.22-4.68 × 104 km2 in 2070, with southern Henan and most Jiangsu being unsuitable for P. vittata growth. Among the 11 environmental variables, the minimum temperature of coldest month (bio6) and temperature annual range (bio7) are the two key factors limiting P. vittata distribution. At bio6 <-5 °C and/or bio7 >33 °C, the regions are unsuitable for P. vittata growth. Based on the MaxEnt model, precipitation had limited effects, so P. vittata can probably survive under both dry and moist environments. This study helps guide phytoremediation of As-polluted soils using P. vittata and provides an example to evaluate habitat suitability of hyperaccumulators at international scales.

Novel PvACR3;2 and PvACR3;3 genes from arsenic-hyperaccumulator Pteris vittata and their roles in manipulating plant arsenic accumulation

Arsenite (AsIII) antiporter ACR3 is crucial for arsenic (As) translocation and sequestration in As-hyperaccumulator Pteris vittata, which has potential for phytoremediation of As-contaminated soils. In this study, two new ACR3 genes PvACR3;2 and PvACR3;3 were cloned from P. vittata and studied in model organism yeast (Saccharomyces cerevisiae) and model plant tobacco (Nicotiana tabacum). Both ACR3s mediated AsIII efflux in yeast, decreasing its As accumulation and enhancing its As tolerance. In addition, PvACR3;2 and PvACR3;3 were expressed in tobacco plant. Localized on the plasma membrane, PvACR3;2 mediated both AsIII translocation to the shoots and AsIII efflux from the roots in tobacco, resulting in 203 − 258% increase in shoot As after exposing to 5 μM AsIII under hydroponics. In comparison, localized to the vacuolar membrane, PvACR3;3 sequestrated AsIII in tobacco root vacuoles, leading to 18 − 20% higher As in the roots and 15 − 36% lower As in the shoots. Further, based on qRT-PCR, both genes were mainly expressed in P. vittata fronds, indicating PvACR3;2 and PvACR3;3 may play roles in AsIII translocation and sequestration in the fronds. This study provides not only new insights into the functions of new ACR3 genes in P. vittata, but also important gene resources for manipulating As accumulation in plants for phytoremediation and food safety.

Remediation of arsenic-contaminated paddy soil by intercropping aquatic vegetables and rice

Soil contamination by arsenic (As) is an important environmental issue globally. Intercropping of hyperaccumulators with main crop is typically applied for remediation of As-contaminated soil. Most hyperaccumulators are wild plants with small biomass and slow growth rates. Thus, remediation is slow. Here, we propose an effective intercropping system for remediation of As-contaminated paddy soil. Four treatments—intercropping with water spinach (Ipomoea aquatica Forsk) (T1), water celery (Oenanthe javanica (Blume) DC.) (T2), or Guangdong white arrowhead (Sagittaria sagittifolia L. var) (T3), with rice (Oryza sativa L.) monoculture (control, CK)—were used. Compared with the CK, grain yield per plant of rice under T1 and T2 increased by 58.13% and 10.48%, respectively, but decreased by 46.90% in T3. As concentration, bioaccumulation factor, and translocation factor in brown rice were significantly lower in the intercropping treatments than in CK. As removal by water spinach was 7.04 and 1.47 times that by water celery and arrowhead, respectively. The pH of paddy soil was significantly higher in all treatments than in CK, and iron plaque on rice roots under T1 and T2 decreased significantly but increased significantly under T3 compared with that of CK. Rice intercropped with water spinach had the best remediation effect. Novelty Statement: We believe that the following highlights of this manuscript will make it interesting to general readers of this journal. First, in recent years, many articles about intercropping system for the remediation of soil heavy metal pollution focus on dry land, and few studies have focused on paddy soil. The present study was on arsenic-contaminated paddy soil remediation. Second, water spinach, water celery, and arrowhead have great potential for phytoremediation. Studies have shown that these three aquatic vegetables play a role in the removal of certain pollutants, such as heavy metals. Moreover, when intercropped with rice, they can effectively increase rice yield and reduce rice diseases and insect pests. However, studies on remediation of arsenic-contaminated soil by intercropping aquatic vegetables and rice have not been reported. We propose here a rice-aquatic vegetables (water spinach, water celery and arrowhead) intercropping pattern for remediation of arsenic in soil. Third, according to the arsenic concentration and removal rate, we used a bioaccumulation factor, translocation factor, and arsenic removal per unit area of plants for the quantitative evaluation of the remediation effects of the intercropping systems. We found that the intercropping of rice and water spinach could be used to remediate arsenic-contaminated soil. Moreover, the extraction contents of arsenic using intercropping with water spinach was higher than that achieved in a previous study that applied intercropping with the arsenic hyperaccumulator Pteris vittata over the same growth time. This study provides a reference for realizing both remediation and increased production in arsenic-contaminated soil and for promoting sustainable development of agriculture.

Efficient arsenate reduction in As-hyperaccumulator Pteris vittata are mediated by novel arsenate reductases PvHAC1 and PvHAC2

Arsenic-hyperaccumulator Pteris vittata is efficient in As absorption, reduction, and translocation. But the molecular mechanisms and locations of arsenate (AsV) reduction in P. vittata are still unclear. Here, we identified two new arsenate reductase genes from P. vittata, PvHAC1 and PvHAC2. Two PvHAC genes encoded a rhodanase-like protein, which were localized in the cytoplasm and nucleus. Both recombinant Escherichia coli strains and transgenic Arabidopsis thaliana lines showed arsenate reductase ability after expressing PvHAC genes. Further, expressing PvHAC2 enhanced As tolerance and reduced As accumulation in A. thaliana shoots under AsV exposure. Based on expression pattern analysis, PvHAC1 and PvHAC2 were predominantly expressed in the rhizomes and fronds of P. vittata. Different from those of HAC homologous genes in non-hyperaccumulators, little PvHAC was expressed in the roots. Besides, PvHAC1 expression was strongly upregulated under AsV exposure but not AsIII. The data suggest that arsenate reductase PvHAC1 in the rhizomes coupled with arsenate reductase PvHAC2 in the fronds of P. vittata played a critical role in As-hyperaccumulation by P. vittata, which helps to further improve its utility in phytoremediation of As-contaminated soils.

Expression of New Pteris vittata Phosphate Transporter PvPht1;4 Reduces Arsenic Translocation from the Roots to Shoots in Tobacco Plants.

Arsenic-hyperaccumulator Pteris vittata is efficient in As uptake, probably through phosphate transporters (Pht). Here, for the first time, we cloned a new PvPht1;4 gene from P. vittata and investigated its role in arsenate (AsV) uptake and transport in yeast and transgenic tobacco plants. On the basis of quantitative real-time polymerase chain reaction (qRT-PCR), PvPht1;4 was abundantly expressed in P. vittata fronds and roots, with its transcripts in the roots being induced by both P deficiency and As exposure. PvPht1;4 was localized to the plasma membrane, which complemented a yeast-mutant defective in P uptake and showed higher P transport affinity than PvPht1;3. Under AsV exposure, PvPht1;4 yeast transformants showed comparable tolerance as PvPht1;3, but higher As accumulation than PvPht1;2 transformants, indicating that PvPht1;4 had considerable AsV and P transport activity. However, in soil and hydroponic experiments, PvPht1;4 expressing tobacco lines accumulated 26-44 and 37-55% lower As in the shoots than wild type plants, with lower root-to-shoot As translocation. In the roots of PvPht1;4 lines, higher glutathione (GSH) contents and expression levels of GSH synthetase gene NtGSH2 were observed. In addition, the transcripts of AsIII-GSH transporter NtABCC1 in PvPht1;4 lines were upregulated. The data suggested that PvPht1;4 lines probably detoxified As by reducing AsV to AsIII, which was then complexed with GSH and stored in the root vacuoles, thereby reducing As translocation in transgenic tobacco. Given its strong AsV transport capacity, expression of PvPht1;4 provides a new molecular approach to reduce As accumulation in plant shoots.

Arsenic accumulation and distribution in Pteris vittata fronds of different maturity: Impacts of soil As concentrations

Arsenic (As) hyperaccumulator Pteris vittata is efficient in As uptake, translocation and accumulation, but the impacts of soil As concentrations on As accumulation and distribution in P. vittata are still unclear. The impacts of soil As (7.3, 63 and 228 mg kg−1) on plant growth and As accumulation in P. vittata after 6 months of growth were evaluated. Arsenic concentrations in the roots, midribs and pinna margin of P. vittata fronds of different maturity were determined by inductively coupled plasma mass spectrometry (ICP-MS) and scanning electron microscopy coupled with an energy dispersive spectrometer (SEM-EDS). While moderate As level at As63 didn’t impact P. vittata growth, higher As at As228 decreased plant biomass by 38%. Under As stress, more As was accumulated in the senescing fronds (47%) and mature fronds (11%) than the young fronds. In senescing fronds, As concentrations in pinna margin were 2.3 times that of the midribs, consistent with As-induced necrotic symptom. Arsenic distribution based on SEM-EDS analysis revealed good correlation between Si and As in the pinnae (r = 0.49). Our data showed that As accumulation in pinna margin caused necrosis and Si may have a potential role in As detoxification in P. vittata.

Heterologous Expression of Pteris vittata Phosphate Transporter PvPht1;3 Enhances Arsenic Translocation to and Accumulation in Tobacco Shoots.

Arsenic-hyperaccumulator Pteris vittata is efficient in As accumulation and has been used in phytoremediation of As-contaminated soils. Arsenate (AsV) is the predominant As species in aerobic soils and is taken up by plants via phosphate transporters (Pht) including P. vittata. In this work, we cloned the PvPht1;3 full length coding sequence from P. vittata and investigated its role in As accumulation by yeast and plants. PvPht1;3 complemented a yeast P uptake mutant strain and showed a stronger affinity and transport capacity to AsV than PvPht1;2. In transgenic tobacco, PvPht1;3 enhanced AsV absorption and translocation, increasing As accumulation in the shoots under both hydroponic and soil experiments. On the basis of the expression patterns via qRT-PCR, PvPht1;3 was strongly induced by P deficiency but not As exposure. To further understand its expression pattern, transgenic Arabidopsis thaliana and soybean expressing the GUS reporter gene, driven by PvPht1;3 promoter, were produced. The GUS staining showed that the reporter gene was mainly expressed in the stele cells, indicating that PvPht1;3 was expressed in stele cells and was likely involved in P/As translocation. Taken together, the data suggested that PvPht1;3 was a high-affinity AsV transporter and was probably responsible for efficient As translocation in P. vittata. Our results suggest that expressing PvPht1;3 enhances As translocation and accumulation in plants, thereby improving phytoremediation of As-contaminated soils.

Chelator-assisted phytoextraction of arsenic, cadmium and lead by Pteris vittata L. and soil microbial community structure response

Using biodegradable chelators to assist in phytoextraction may be an effective approach to enhance the heavy-metal remediation efficiencies of plants. A pot experiment was conducted to investigate the effects of ethylenediamine disuccinic acid (EDDS), citric acid (CA), and oxalic acid (OA) on the growth of the arsenic (As) hyperaccumulator Pteris vittata L., its arsenic (As), cadmium (Cd), and lead (Pb) uptake and accumulation, and soil microbial responses in multi-metal(loid)-contaminated soil. The addition of 2.5-mmol kg−1 OA (OA-2.5) produced 26.7 and 14.9% more rhizoid and shoot biomass, respectively compared with the control, while EDDS and CA treatments significantly inhibited plant growth. The As accumulation in plants after the OA-2.5 treatment increased by 44.2% and the Cd and Pb accumulation in plants after a 1-mmol kg−1 EDDS treatment increased by 24.5 and 19.6%, respectively. Soil urease enzyme activities in OA-2.5 treatment were significantly greater than those in the control and other chelator treatments (p < 0.05). A PCR–denatured gradient gel electrophoresis analysis revealed that with the addition of EDDS, CA and OA enhanced soil microbial diversity. It was concluded that the addition of OA-2.5 was suitable for facilitating phytoremediation of soil As and did not have negative effects on the microbial community.

Phytate promoted arsenic uptake and growth in arsenic-hyperaccumulator Pteris vittata by upregulating phosphorus transporters

While phosphate (P) inhibits arsenic (As) uptake by plants, phytate increases As uptake by As-hyperaccumulator Pteris vittata. Here we tried to understand the underling mechanisms by investigating the roles of phytate in soil As desorption, P transport in P. vittata, short-term As uptake, and plant growth and As accumulation from soils. Sterile soil was used to exclude microbial degradation on phytate. Results showed that inorganic P released 3.3-fold more As than that of phytate from soil. However, P. vittata accumulated 2–2.5 fold more As from soils with phytate than that in control and P treatment. In addition, different from P suppression on As uptake, solution uptake experiment showed that As uptake in phytate treatment was comparable to that of control under 0.1–7.5 μM As after 1–24 h. Moreover, responding to phytate, P. vittata P transporter PvPht1;3 increased by 3-fold while PvPht1;1 decreased by 65%. The data suggested that phytate upregulated PvPht1;3, thereby contributing to As uptake in P. vittata. Our results showed that, though with lower As release from soil compared to P, phytate induced more As uptake and better growth in P. vittata by upregulating P transporters.

The Soil Amendments to Improve the Efficiency of the Intercropping System of Pteris vittata and Morus alba

Intercropping of arsenic (As) hyperaccumulator Pteris vittata and cash crop Morus alba could improve the As concentration in the hyperaccumulator but decrease As concentration in the intercropped crop. The effects of several amendments on the transfer of As were investigated to determine an enhancement strategy for the intercropping system of P. vittata and M. alba. Phosphorus, in the form of Ca(H2PO4)2, promoted the release of As to the soil solution and apparently increased the As removal from the soil by 42% compared with the untreated variant. The addition of FeSO4 and CaCO3 decreased As concentration in the soil solution and the uptake of As by both plant species. The As levels in the mulberry leaves remained under the threshold limits of feedstuffs in China. Intercropping was confirmed as an applicable strategy to manage contaminated soil. Hence, under the condition that all treatments produced safe mulberry leaves, Ca(H2PO4)2 was the appropriate amendment to achieve the highest As removal rate, whereas FeSO4 could lower the risk of As to further migrate to another medium.

Phosphate Transporter PvPht1;2 Enhances Phosphorus Accumulation and Plant Growth without Impacting Arsenic Uptake in Plants.

Phosphorus is an important macronutrient for plant growth and is acquired by plants mainly as phosphate (P). Phosphate transporters (Phts) are responsible for P and arsenate (AsV) uptake in plants including arsenic-hyperaccumulator Pteris vittata. P. vittata is efficient in AsV uptake and P utilization, but the molecular mechanism of its P uptake is largely unknown. In this study, a P. vittata Pht, PvPht1;2, was cloned and transformed into tobacco ( Nicotiana tabacum). In hydroponic experiments, all transgenic lines displayed markedly higher P content and better growth than wild type, suggesting that PvPht1;2 mediated P uptake in plants. In addition, expressing PvPht1;2 also increased the shoot/root 32P ratio by 69-92% and enhanced xylem sap P by 46-62%, indicating that PvPht1;2 also mediated P translocation in plants. Unlike many Phts permeable to AsV, PvPht1;2 showed little ability to transport AsV. In soil experiments, PvPht1;2 also significantly increased shoot biomass without elevating As accumulation in PvPht1;2 transgenic tobacco. Taken together, our results demonstrated that PvPht1;2 is a specific P transporter responsible for P acquisition and translocation in plants. We envisioned that PvPht1;2 can enhance crop P acquisition without impacting AsV uptake, thereby increasing crop production without compromising food safety.

Intercropping efficiency of four arsenic hyperaccumulator Pteris vittata populations as intercrops with Morus alba.

Soils that are slightly or moderately contaminated with arsenic (As) can be safely utilized by intercropping As hyperaccumulator Pteris vittata with cash crops. Introducing hyperaccumulators into crop planting systems results in the alleviation of the adverse effects of As and competition effect for resources. The balance between these two effects determines intercropping efficiency. The effect of using different hyperaccumulator populations on such balance is the focus of this study. Through a tank experiment, four P. vittata populations were compared on the basis of their intercropping efficiencies and physiological and morphological characteristics. The evaluation of the intercropping efficiency of P. vittata was mainly based on the capabilities of the species to promote growth and decrease As concentrations in intercropped Morus alba. Two populations of P. vittata were appropriate for intercropping with M. alba, with the alleviation effect of As harm as the main effect on the intercropping system. These populations showed extensive root overlap with M. alba and efficient uptake of bioavailable As, thus depleting As in the rhizosphere and lowering As risk. After different P. vittata populations were used, varied interspecific interactions were observed. Root overlap and aboveground morphological parameters are the key factors determining intercropping efficiency among P. vittata populations.

Phytate induced arsenic uptake and plant growth in arsenic-hyperaccumulator Pteris vittata

Phytate is abundant in soils, which is stable and unavailable for plant uptake. However, it occurs in root exudates of As-hyperaccumulator Pteris vittata (PV). To elucidate its effect on As uptake and growth, P. vittata were grown on agar media (63 μM P) containing 50 μM As and/or 50 or 500 μM phytate with non As-hyperaccumulator Pteris ensiformis (PE) as a congeneric control for 60 d. Phytate induced efficient As and P uptake, and enhanced growth in PV, but had little effects on PE. The As concentrations in PV fronds and roots were 157 and 31 mg kg−1 in As50+phytate50, 2.2- and 3.1-fold that of As50 treatment. Phosphorus uptake by PV was reduced by 27% in As treatment than the control (P vs. P+As) but increased by 73% comparing phytate500 to phytate500+As, indicating that PV effectively took up P from phytate. Neither As nor phytate affected Fe accumulation in PV, but phytate reduced root Fe concentration in PE (46–56%). As such, the increased As and P and the unsuppressed Fe uptake in PV probably promoted PV growth. Thus, supplying phytate to As-contaminated soils may promote As uptake and growth in PV and its phytoremediation ability.

Arsenic-hyperaccumulator Pteris vittata efficiently solubilized phosphate rock to sustain plant growth and As uptake

Phosphorus (P) is one of the most important nutrients for phytoremediation of arsenic (As)-contaminated soils. In this study, we demonstrated that As-hyperaccumulator Pteris vittata was efficient in acquiring P from insoluble phosphate rock (PR). When supplemented with PR as the sole P source in hydroponic systems, P. vittata accumulated 49% and 28% higher P in the roots and fronds than the −P treatment. In contrast, non-hyperaccumulator Pteris ensiformis was unable to solubilize P from PR. To gain insights into PR solubilization by plants, organic acids in plant root exudates were analyzed by HPLC. The results showed that phytic acid was the predominant (>90%) organic acid in P. vittata root exudates whereas only oxalic acid was detected in P. ensiformis. Moreover, P. vittata secreted more phytic acid in −P and PR treatments. Compared to oxalic acid, phytic acid was more effective in solubilizing PR, suggesting that phytic acid was critical for PR utilization. Besides, secretion of phytic acid by P. vittata was not inhibited by arsenate. Our data indicated that phytic acid played an important role in efficient use of insoluble PR by P. vittata, shedding light on using insoluble PR to enhance phytoremediation of As-contaminated soils.

Microbial siderophores and root exudates enhanced goethite dissolution and Fe/As uptake by As-hyperaccumulator Pteris vittata

Arsenic (As) in soils is often adsorbed on Fe-(hydro)oxides surface, rendering them more resistant to dissolution, which is undesirable for phytoremediation of As-contaminated soils. Arsenic hyperaccumulator Pteris vittata prefers to grow in calcareous soils where available Fe and As are low. To elucidate its mechanisms of acquiring Fe and As from insoluble sources in soils, we investigated dissolution of goethite with pre-adsorbed arsenate (AsV; As-goethite) in presence of four organic ligands, including two root exudates (oxalate and phytate, dominant in P. vittata) and two microbial siderophores (PG12-siderophore and desferrioxamine B). Their presence increased As solubilization from As-goethite from 0.03 to 0.27–5.33 mg L−1 compared to the control. The siderophore/phytate bi-ligand treatment released 7.42 mg L−1 soluble Fe, which was 1.2-fold that of the sum of siderophore and phytate, showing a synergy in promoting As-goethite dissolution. In the ligand-mineral-plant system, siderophore/phytate was most effective in releasing As and Fe from As-goethite. Moreover, the continuous plant uptake induced more As-goethite dissolution. The continued release of As and Fe significantly enhanced their plant uptake (from 0.01 to 0.43 mg plant−1 As and 2.7–14.8 mg plant−1 Fe) and plant growth (from 1.2 to 3.1 g plant−1 fw) in P. vittata. Since microbial siderophores and root exudates often coexist in soil rhizosphere, their synergy in enhancing dissolution of insoluble As-Fe minerals may play an important role in efficient phytoremediation of As-contaminated soils.