Arsenic hyperaccumulation in the Chinese brake fern (Pteris vittata) deters grasshopper (Schistocerca americana) herbivory

Brake fern, Pteris vittata, not only tolerates arsenic but also hyperaccumulates it in the frond. The hypothesis that arsenic hyperaccumulation in this fern could function as a defense against insect herbivory was tested. Fronds from control and arsenic-treated ferns were presented to nymphs of the grasshopper Schistocerca americana. Feeding damage was recorded by visual observation and quantification of the fresh weight of frond left uneaten and number of fecal pellets produced over a 2-d period. Grasshopper weight was determined before and after 5 d of feeding. Grasshoppers consumed significantly greater amounts of the frond tissue, produced more fecal pellets and had increased body weight on control plants compared with grasshoppers fed arsenic-treated ferns. Very little or none of the arsenic-treated ferns were consumed indicating feeding deterrence. In a feeding deterrent experiment with lettuce, sodium arsenite at 1.0 mm deterred grasshoppers from feeding whereas 0.1 mm did not. In a choice experiment, grasshoppers preferred to feed on lettuce dipped in water compared with lettuce dipped in 1.0 mm sodium arsenite. Our results show that arsenic hyperaccumulation in brake fern is an elemental defense against grasshopper herbivory.     

蜈蚣草的解剖结构与组织化学特征

蜈蚣草（Pteris vittata）是多年生的超积累砷植物,并用于修复受重金属污染的土壤。利用光学显微镜和荧光显微镜来研究蜈蚣草的解剖结构及组织化学特征,以此明确该物种适应干旱岩生环境,以及具有离子超富集作用的特点。结果表明（：1）蜈蚣草孢子体的根状茎、不定根和叶的结构均为初生结构,不定根的结构由内而外包括维管柱、内皮层、皮层、木质化厚壁组织层和表皮。（2）根状茎结构由内而外包括网状中柱、内皮层、皮层、表皮外覆盖的角质层。（3）羽状复叶的总叶柄的结构由内而外包括维管束、内皮层、皮层、厚壁组织层、表皮外覆盖的角质层。叶片为异面叶,表皮内方具厚壁层,叶表皮具角质层,仅下表皮有气孔。（4）蜈蚣草根表皮、皮层与根毛的表面富含果胶,皮层木质化;黄连素离子通透性试验结果显示,根毛、根表皮和皮层滞留大量黄连素离子。综上,植物体的内皮层、木质化厚壁组织层、异面叶和厚的角质层结构说明蜈蚣草适应岩生环境,根具木质化皮层和富含果胶的组织化学特点,以及离子通透性试验表明其与离子超积累功能有关。     

An arsenic hyperaccumulating fern,Pteris vittata L. (Pteridaceae) broadly affects terrestrial invertebrate abundance

1. The Chinese brake fern (Pteris vittata L.; Pteridaceae) can accumulate up to 27 000 mg kg^?1 dry wt. of arsenic (As) from the soil into its above‐ground biomass. They may use this As to deter invertebrate threats. 2. This study explored how As concentrations [As] in the fern, and in soil associated with the fern, influenced the abundance and composition of various invertebrates. 3. Populations of P. vittata were identified in the field. Soils from the base of the fern and from 3 m away of each plant were collected and pitfall traps were installed. Soil and fern arsenic concentrations ([As]) were measured via inductively coupled plasma mass spectrometry and invertebrates were identified to order and classified by feeding guild. 4. Increased [As] did not affect all feeding guilds and orders equally. For example, individual herbivore abundance did not decrease as [As] increased, but predator abundance did. In many cases, the impact of soil [As] on invertebrates depended on the distance from the fern. Fern [As] also influenced components of the community, but only at 3 m away from the fern. Furthermore, the abundances of many invertebrate groups were higher beneath the fern, where [As] was higher. 5. These results suggest that hyperaccumulated As can impact the invertebrate community, but the defensive benefits of hyperaccumulation are more complex than have been previously described. The authors advocate that future studies examining the potential defensive benefits of hyperaccumulation should do so in a natural setting that incorporates this complexity and invertebrate richness.     

Exploring the pattern of phenotypic and genetic polymorphism in the arsenic hyperaccumulator Pteris vittata L. (Chinese brake fern)

Background and aims

The trace element uptake process of plants is a key factor in assessing the risks of trace element build-up in agricultural soils. Scarce information exists on the trace element dynamic uptake of plants grown in the field, especially on those potentially hazardous. In this study, the uptake process of As, Cd, Cu, and Zn in maize plants was quantified and characterized throughout the entire season.

Methods

Along two seasons, the uptake dynamics of field-grown maize plants in absorbing the soil borne trace elements was examined. Biomass production and the concentration of the elements in plant and soil solution samples were determined. A kinetic model was employed to characterize the uptake by plants.

Results

The kinetic parameters of the uptake process, maximum cumulative uptake rate, U _max , time to reach 50 % of U _max , t _U50 , and reciprocal of the uptake rate, b _U when followed throughout the season in terms of the plant’s growing degree days remained constant between seasons and were element specific. In spite of the large amount biomass produced, maize plants extracted minute quantities of Cd and As. Increasing cumulative uptake rates of As, Cd, Cu, and Zn from the soil took place primarily in the early half of the growing season when the biomass accumulation was still less than 50 % of the maximum harvested biomass. The element-specific plant uptake factor (PUF), which denote the partition of trace elements between the soil solution and plant phases, decreased following a first-order kinetics along the growing period, did not show any significant difference between seasons, and, at maturity stage, followed the sequence Cd≥Zn>Cu≥As.

Conclusions

The uptake process of the elements was adequately described by the kinetic model, showing similar patterns but different magnitude and distribution in the plant. The extraction of Cd and As by plants is low in comparison to common inputs through fertilizer applications into maize production systems, indicating potential risk of trace element accumulation in soils. The PUF may be estimated according to the kinetics parameters of the uptake process. On a per-unit-soil solution element basis, Cd and Zn would be more susceptible to the soil-to-plant transfer than As and Cu.     

Molecular phylogeny and biogeography of the fern genus Pteris (Pteridaceae)

Background and Aims

Pteris (Pteridaceae), comprising over 250 species, had been thought to be a monophyletic genus until the three monotypic genera Neurocallis, Ochropteris and Platyzoma were included. However, the relationships between the type species of the genus Pteris, P. longifolia, and other species are still unknown. Furthermore, several infrageneric morphological classifications have been proposed, but are debated. To date, no worldwide phylogenetic hypothesis has been proposed for the genus, and no comprehensive biogeographical history of Pteris, crucial to understanding its cosmopolitan distribution, has been presented.

Methods

A molecular phylogeny of Pteris is presented for 135 species, based on cpDNA rbcL and matK and using maximum parsimony, maximum likelihood and Bayesian inference approaches. The inferred phylogeny was used to assess the biogeographical history of Pteris and to reconstruct the evolution of one ecological and four morphological characters commonly used for infrageneric classifications.

Key Results

The monophyly of Pteris remains uncertain, especially regarding the relationship of Pteris with Actiniopteris + Onychium and Platyzoma. Pteris comprises 11 clades supported by combinations of ecological and morphological character states, but none of the characters used in previous classifications were found to be exclusive synapomorphies. The results indicate that Pteris diversified around 47 million years ago, and when species colonized new geographical areas they generated new lineages, which are associated with morphological character transitions.

Conclusions

This first phylogeny of Pteris on a global scale and including more than half of the diversity of the genus should contribute to a new, more reliable infrageneric classification of Pteris, based not only on a few morphological characters but also on ecological traits and geographical distribution. The inferred biogeographical history highlights long-distance dispersal as a major process shaping the worldwide distribution of the species. Colonization of different niches was followed by subsequent morphological diversification. Dispersal events followed by allopatric and parapatric speciation contribute to the species diversity of Pteris.     

Root distributions and elemental accumulations of Chinese brake (Pteris vittata L.) from As-contaminated soils

A field investigation was conducted to understand the root distributions and elemental accumulations of Chinese brake (Pteris vittata L.), an As-hyperaccumulator, grown in soils with a gradient of As concentration near an arsenic sulphide mine. The root distribution was affected not only by the levels of soil As, but also by soil texture. Plants grew better in sandy loam soils than in clay soils. Increases in the ratio of frond biomass to underground biomass were correlated with decreasing soil As concentration. Root densities of the plant decreased from 0–10 cm, 10–20 cm to 20–30 cm in the soil profiles. Most of the roots were concentrated in the upper 0–10 cm layer. Under high As conditions, As concentrations in different tissues followed the trends: pinnae > rhizomes ≈ roots of 0–10 cm > roots of 10–20 cm > roots of 20–30 cm > petioles, however, As concentrations in pinnae were higher than those in rhizomes under low As conditions. The rhizomes and pinnae were the main As pools, storing 75–86% of the total As uptaken by the plants. The rhizome, a `buffer-storage' for plant As, maintained high concentrations of As under high soil As while the pinnae became the most important organ of storing the As under low soil As. Chinese brake might possess the ability of adjusting its As-storage under different soil As levels. The plant can not only hyperaccumulate As from the soils, but also enriched P and Ni from the soils and translocated them to the fronds. It is important to improve the root distribution for phytoremediation of As-contaminated soils using Chinese brake.     

Phytoremediation of arsenic and lead in contaminated soil using Chinese brake ferns (Pteris vittata) and Indian mustard (Brassica juncea)

Field and greenhouse experiments were performed to assess the performance of phytoremediation of arsenic and lead from contaminated soil at an EPA Superfund site (Barber Orchard). Chinese Brake ferns (Pteris vittata) were used to extract arsenic. On average, fern shoot arsenic concentrations were as high as 20 times the soil arsenic concentrations under field conditions. It was estimated that 8 years would be required to reduce the acid-extractable portion of soil arsenic to safe levels (40 mg/kg). The effect of soil pH on arsenic extraction was also investigated. Results indicate that increasing soil pH may improve arsenic removal. Indian mustard plants (Brassica juncea) were used under greenhouse conditions to phytoextract soil lead. EDTA was applied to soil and was found to improve lead extraction. When the EDTA concentration was 10 mmol EDTA/kg soil in soil containing 338 mg Pb/kg soil, mustard plants extracted approximately 32 mg of lead. In conclusion, phytoremediation would be a suitable alternative to conventional remediation techniques, especially for soils that do not require immediate remediation.     

A new species of fern of the genus Pteris (Filicales: Pteridaceae) endemic to Costa Rica

The new fern species Pteris herrerae A. Rojas & M. Palacios, endemic to Costa Rica, is described. It differs from P. decurrens C. Presl in basal segments reduced to 1/5-1/2 of the next segment (vs. 2/3-3/4), basal pinnae not bifurcated (vs. bifurcated), pinnae apex mucronate (vs. acuminate) and segment apex undulate (vs. dentate). It differs from Pteris consanguinea in the elliptic pinnae (vs. oblong), two segments reduced on the base (vs. lack), segments entire to undulate (vs. dentate), basal pinnae without basiscopic lobes (vs. with basiscopic lobes) and segment apex entire to undulate (vs. dentate).     

Phytoremediation of arsenic-contaminated groundwater by the arsenic hyperaccumulating fern Pteris vittata L

Arsenic concentrations in a much larger fraction of U.S. groundwater sources will exceed the maximum contaminant limit when the new 10 microg L(-1) EPA standard for drinking water takes effect in 2006. Thus, it is important to develop remediation technologies that can meet this new standard. Phytoremediation of arsenic-contaminated groundwater is a relatively new idea. In this research, an arsenic-hyperaccumulating fern, commonly known as Chinese Brake fern (Pteris vittata L.), was grown hydroponically to examine its effectiveness in arsenic removal from what is believed to be herbicide-contaminated groundwater. One plant grown in 600 mL of groundwater effectively reduced the arsenic concentration from 46 to less than 10 microg L(-1) in 3 days. Re-used plants continued to take up arsenic from the groundwater, albeit at a slower rate (from 46 to 20 microg L(-1) during the same time). Young fern plants were more efficient in removing arsenic than were older fern plants of similar size. The addition of a supplement of phosphate-free Hoagland nutrition to the groundwater had little effect on arsenic removal, but the addition of phosphate nutrition significantly reduced its arsenic affinity and, thus, inhibited the arsenic removal. This study suggested that Chinese Brake has some potential to remove arsenic from groundwater.     

刈割对蜈蚣草的砷吸收和植物修复效率的影响

以野生苗移栽的蜈蚣草为试材 ,通过盆栽试验研究了收获次数对蜈蚣草生长、砷吸收和植物修复效率的影响。结果表明 :在 3次收获中 ,随着收获次数的增加 ,不同砷浓度处理之间蜈蚣草生物量的差异逐步缩小 ;不加砷的对照处理中 ,每次收获后的砷吸收速率下降趋势 ,而在 3个加砷处理中 ,第 2次收获和第 3次收获的蜈蚣草的吸砷速率为 6 3～ 75 μg/ (plant· d)、4 4～ 5 5μg/ (plant· d) ,均显著高于第 1次收获时的吸收速率。表明多次收获并没有降低砷的积累速度。由此可见 ,通过适当增加蜈蚣草的收获次数是提高砷修复效率的一种策略     

Sexual Phase of Three Species of Pteris (Pteridaceae)

This work describes and compares morphogenesis of the gametophyte, or sexual phase, of three ferns, Pteris deflexa, P. denticulata and P. tristicula, through a palynological analysis of live spores. This study was conducted under laboratory conditions; observations and pictures were made using light and scanning electron microscopy. The pattern of spore germination corresponds to the Vittaria type and gametophytic development corresponds to the Ceratopteris type. We observed that the polymorphous spores of P. denticulata and P. tristicula are viable. Ultrastructural details of the gametophytes revealed that P. denticulata and P. tristicula form antheridia different from those of the remaining Pteris species; the archegonia have necks formed by 4 rows of 4 cells each. The type of reproduction is sexual and sporophytes arise 90 days after sowing.     

Cadmium uptake,localization and stress-induced morphogenic response in the fern Pteris vittata

Cadmium uptake, tissue localization and structural changes induced at cellular level are essential to understand Cd tolerance in plants. In this study we have exposed plants of Pteris vittata to different concentrations of CdCl₂ (0, 30, 60, 100 μM) to evaluate the tolerance of the fern to cadmium. Cadmium content determination and its histochemical localization showed that P. vittata not only takes up, but also transports and accumulates cadmium in the aboveground tissues, delocalizing it mainly in the less bioactive tissues of the frond, the trichomes and the scales. Cadmium tolerance in P. vittata was strictly related to morphogenic response induced by the metal itself in the root system. Adaptive response regarded changes of the root apex size, the developmental pattern of root hairs, the differentiation of xylem elements and endodermal suberin lamellae. All the considered parameters suggest that, in our experimental conditions, 60 μM of Cd may represent the highest concentration that P. vittata can tolerate; indeed this Cd level even improves the absorbance features of the root and allows good transport and accumulation of the metal in the fronds. The results of this study can provide useful information for phytoremediation strategies of soils contaminated by Cd, exploiting the established ability of P. vittata to transport, delocalize in the aboveground biomass and accumulate polluting metals.     

Antheridiogenesis in the fern Pteris vittata. I. Photocontrol of antheridium formation

When spores of the fern Pteris vittata are induced to germinate on water in the light, they develop solely a rhizoid. Upon addition of mineral salt medium and transfer into darkness they develop male gametangia directly at the spore. This precocious antheridiogenesis is inhibited by light. Ohytochrome and a blue-light photoreceptor are both involved in the control. Ions are required for the light-dependent inhibition of antheridia formation. It is concluded that spores are determined generatively while light inhibits the generative tendency and promotes the vegetative development.     

Chemical constituents from Pteris ensiformis Burm. (Pteridaceae)

Phytochemical study of Pteris ensiformis Burm. resulted in the isolation of eleven compounds, including three sesquiterpenoids (1–3), one alkaloid (5), one amino acid (6), one flavonoid (7), four phenolic acids (4, 8, 9 and 10), and one coumarin (11). Structures of these compounds were deduced from the ESI-MS and NMR data. This is the first report on the isolation of compounds 1, 2, 7 and 11 from P. ensiformis, compounds 6 and 9 from the genus Pteris, and compound 5 from the family Pteridaceae. In addition, a detailed discussion of the chemotaxonomic significance of these compounds was presented.     

Light-induced synchrony of cell division in the protonema of the fern,Pteris vittata

Summary In protonemata of Pteris vittata grown for 6 days under red light, which brings about a marked depression of mitotic activity, the first division of the cells was synchronously induced by irradiation with blue light, and subsequent cell divisions were also promoted. The peak of the mitotic index reached a maximum of about 70% at 11.5 hrs, and 90% of all protonemata divided between the 11th and 13th hour after exposure to blue light. When the protonemata were continuously irradiated with blue light, synchronism of the next cell division in the apical cells decreased to a mitotic index of about 30%, and further divisions occurred randomly.The synchronization of cell division was found to be a combined effect of red and blue light. Red light maintained the cells in the early G₁ phase of the cell cycle; blue light caused the cells to progress synchronously through the cell cycle, with an average duration of 12 hr. By using ³H-thymidine, the average duration of the G₁, S, G₂ and M phases was determined to be about 3.5, 5, 2.5 and 1 hr, respectively.Synchronous cell division could be induced in older protonemata grown for 6 to 12 days in red light and even in protonemata having two cells. It could be repeated in the same protonema by reexposure to red light for 24 hrs or more before another irradiation with blue light.     

Chemical constituents of Pteris wallichiana J.Agardh (Pteridaceae)

Chemical investigation on the whole plants of Pteris wallichiana J. Agardh led to the isolation and identification of a new pterosin glycoside, ptemuloside A (1), along with other seventeen known compounds (2–18) (Fig. 1). The structure of compound 1 was elucidated by means of spectroscopic techniques, including HRESIMS, and 1D and 2D NMR experiments. Compounds 2, 3, and 5–17 are first time reported from the title plant. The pterosins in P. wallichiana could serve as its chemotaxonomic markers.     

Arsenic-resistant proteobacterium from the phyllosphere of arsenic-hyperaccumulating fern (Pteris vittata L.) reduces arsenate to arsenite

An arsenic-resistant bacterium, AsRB1, was isolated from the fronds of Pteris vittata grown in a site contaminated with copper chromium arsenate. The bacterium exhibited resistance to arsenate, arsenite, and antimony in the culture medium. AsRB1, like Pseudomonas putida, grew on MacConkey and xylose-lactose-desoxycholate agars and utilized citrate but, unlike P. putida, was positive for indole test and negative for oxidase test. A phylogenetic analysis of the 16S rRNA gene showed that AsRB1 is a proteobacterium of the beta subclass, related to Pseudomonas saccharophila and Variovorax paradoxus. Following an exogenous supply of arsenate, most arsenic occurred as arsenite in the medium and the cell extracts, suggesting reduction and extrusion of arsenic as the mechanism for arsenic resistance in AsRB1.     

New species in Adiantum and Pteris (Pteridaceae) from the Andes

We describe, discuss, and illustrate four new species, one inAdiantum (A. capillatum A. R. Sm. &; Prado, from Ecuador) and three species inPteris (P. lellingeri A. R. Sm. &; Prado, from Bolivia;P. venezuelensis A. R. Sm. &; Prado, from Venezuela; andP. websteri A. R. Sm. &; Prado, from Ecuador, Colombia, and Bolivia).