Phytoextraction: simulating uptake and translocation of arsenic in a soil-plant system

The uptake, transport, and accumulation of metals by plants are functions central to successful phytoextraction. This study investigates the uptake and translocation of arsenic from a contaminated sandy soil by a mature Chinese brake fern (Pteris vittata L.). An existing mathematical model for the coupled transport of water, heat, and solutes in the soil-plant-atmosphere continuum (CTSPAC) was modified to examine the flow of water as well as the uptake and translocation of total arsenic in the xylem of the fern. This model was calibrated using greenhouse measurements before its application. Simulation results showed that about 20% of the soil arsenic was removed by the fern in 10 d, of which about 90% of the arsenic was stored in the fronds and 10% in the roots. Although arsenic mass in the plant tissues increased consecutively with time, arsenic concentration in the xylem sap of the root tips has a typical diurnal distribution pattern: increasing during the day and decreasing at night, resulting from daily variations of frond surface water transpiration. The largest difference in simulated arsenic concentration in the root tips between the day and night was about 5%. This study also suggests that the use of transpiration stream concentration factor (TSCF), which is defined as the ratio of chemical concentration in the xylem sap to that in the external solution, to evaluate the translocation efficiency of arsenic for the hyperaccumulator Chinese brake fern (Pteris vittata L.) could be limited.     

一种新发现的铜积累植物——密毛蕨

对铜尾矿上生长的密毛蕨(Petridium revolutum)进行了野外调查分析和温室营养液砂培实验.结果表明,密毛蕨所生长的废铜矿土壤中Cu含量平均为2 432 mg·kg^-1 DW,最高达7 554 mg·kg^-1 DW;地上部生物量平均为18.33 g·plant^-1 DW,最高达40.05 g·plant^-1 DW;地上部Cu含量平均为201 mg·kg^-1 DW,最高达567 mg·kg^-1 DW;地下部Cu含量平均为346 mg·kg^-1 DW,最高达1 723 mg kg^-1 DW;密毛蕨对Cu的转移系数平均为0.81,最高达3.88.在营养液砂培的条件下,Cu 7 mg·L^-1处理没有抑制密毛蕨地上部的生长;密毛蕨体内的Cu含量随着介质中Cu浓度的增加而显著增加,但是大部分的Cu积累在地下部.密毛蕨对Cu具有较强的忍耐和较高的积累能力,可作为修复Cu污染土壤的新材料.     

Comparison of root absorption, translocation and tolerance of arsenic in the hyperaccumulator Pteris vittata and the nonhyperaccumulator Pteris tremula

* Several fern species can hyperaccumulate arsenic, although the mechanisms are not fully understood. Here we investigate the roles of root absorption, translocation and tolerance in As hyperaccumulation by comparing the hyperaccumulator Pteris vittata and the nonhyperaccumulator Pteris tremula. * The two species were grown in a pot experiment with 0-500 mg As kg-1 added as arsenate, and in a short-term (8 h) uptake experiment with 5 microM arsenate under phosphorus-sufficient conditions. * In the pot experiment, P. vittata accumulated up to 2500 mg As kg-1 frond d. wt and suffered no phytotoxicity. P. tremula accumulated<100 mg As kg-1 frond d. wt and suffered severe phytotoxicity with additions of >or=25 mg As kg-1. In the short-term uptake experiment, P. vittata had a 2.2-fold higher rate of arsenate uptake than P. tremula, and distributed more As taken up to the fronds (76%) than did P. tremula (9%). * Our results show that enhanced root uptake, efficient root-to-shoot translocation, and a much elevated tolerance through internal detoxification all contribute to As hyperaccumulation in P. vittata.     

Influence of the arbuscular mycorrhizal fungus Glomus mosseae on uptake of arsenate by the As hyperaccumulator fern Pteris vittata L.

We report for the first time some effects of colonization by an arbuscular mycorrhizal (AM) fungus (Glomus mosseae) on the biomass and arsenate uptake of an As hyperaccumulator, Pteris vittata. Two arsenic levels (0 and 300 mg As kg^–1) were applied to an already contaminated soil in pots with two compartments for plant and hyphal growth in a glasshouse experiment. Arsenic application had little or no effect on mycorrhizal colonization, which was about 50% of root length. Mycorrhizal colonization increased frond dry matter yield, lowered the root/frond weight ratio, and decreased frond As concentration by 33–38%. Nevertheless, transfer of As to fronds showed a 43% increase with mycorrhizal colonization at the higher soil As level. Frond As concentrations reached about 1.6 g kg^–1 (dry matter basis) in non-mycorrhizal plants in the As-amended soil. Mycorrhizal colonization elevated root P concentration at both soil As levels and mycorrhizal plants had higher P/As ratios in both fronds and roots than did non-mycorrhizal controls.     

Allelopathic effects of interrupted fern on northern red oak seedlings: Amelioration bySuillus luteus L.: Fr.

Summary The allelopathic effects of interrupted fern frond leachates on ectomycorrhizal (inoculated) and nonmycorrhizal (noninoculated) northern red oak (Quercus rubra L.) seedlings were investigated. Container-grown northern red oak was inoculated with vegetative mycelium ofSuillus luteus L. Fr. following acorn germination. Noninoculated control seedlings were also maintained. Seedlings were grown in a glasshouse under full sunlight or shaded (25% of full sunlight) conditions. Leachate or deionized water solutions were applied to seedlings eleven times over a 91-day period to simulate a rainfall induced transfer of allelopathic chemicals from fern fronds to the soil. Fern frond leachates significantly reduced seedling survival, however, inoculated seedlings showed less mortality. Chromium concentrations of pooled lateral root or leaf tissue were comparatively higher in tissues exhibiting greater mortality. Root biomass was reduced by fern fern frond leachate applications. Seedling biomass was not significantly affected by fungal inoculation. Our results confirm previous documentation of the allelopathic potential of ferns, and suggest that ectomycorrhizal fungi may ameliorate allelopathic effects of ferns on northern red oak seedling survival and growth.     

Interaction of arsenic and phosphate on their uptake and accumulation in Chinese brake fern

Interactive effects of arsenate (As (V)) and phosphate (Pi) were investigated under hydroponic culture. Arsenic concentrations in fronds and roots of Chinese brake fern (Pteris vittata L.) significantly (p < 0.05) increased with increasing As (V), but decreased (p < 0.05) with increasing Pi in nutrient solution. Phosphate uptake was significantly (p < 0.05) inhibited by 1000 micromol L(-1) As (V). Under 100 micromol L(-1) As (V), frond phosphorus (P) increased at 100 and 1000 micromol L(-1) Pi, and root P increased at 250 micromol L(-1) Pi exposures. Arsenic and P concentrations in fronds and roots of Chinese brake fern were negatively correlated (p < 0.05). Arsenate treatments enhanced As and P transport to fronds, while increasing Pi inhibited their transportation, with highest frond P and As (%) obtained under 100 micromol L(-1) treatment. pH values in nutrient solution increased with increasing exposure time, but decreased with increasing Pi levels. Dissolved organic carbon (DOC) contents (dry weight) in nutrient solution decreased with increasing Pi levels, both for treatments with and without As (V). Arsenate at 1000 micromol L(-1) significantly (p < 0.05) increased DOC contents, especially for treatment without Pi. Six organic acids were detected in root exudates of Chinese brake fern, with oxalic and malic acids being most dominant.     

蕨配子体发育及卵发生的显微结构观察

运用显微观察技术对蕨(Pteridium aquilinum var. latiusculum)配子体发育和卵发生进行了研究。结果表明：(1)蕨孢子黄褐色,四面体形,具三裂缝,接种后3～7 d萌发,经丝状体和片状体阶段发育成原叶体,成熟原叶体雌雄异株或同株。(2)蕨颈卵器产生于生长点下方的表面细胞(颈卵器原始细胞),该细胞经2次分裂形成3层细胞,其上层和下层细胞发育为颈卵器壁细胞,中间细胞为初生细胞,它经2次不等分裂产生3个细胞,分别为卵细胞、腹沟细胞和颈沟细胞;刚产生时,此3个细胞紧贴颈卵器壁,细胞质内液泡较多,随着发育,卵细胞和腹沟细胞之间产生了分离腔,但二者通过孔区相连,在卵细胞上表面可观察到卵膜;此后,颈沟细胞和腹沟细胞逐渐退化,颈卵器壁细胞内具有黑色颗粒物质。连续切片观察发现,成熟卵细胞上表面中央具有受精孔。卵发生的细节尚需超微结构的研究。     

A re-evaluation of plants referred to as Pteridiutn herediae (Colmeiro) Love & Kjellqvist

SHEFFIELD, E., WOLF, P. G., HAUFLER, C. H., RANKER, T. A. & JERMY, A. C, 1989. Are-evaluation of plants referred to as Pteridiutn herediae (Colmeiro) Love & Kjellqvist. Based on perceived differences in morphology, chromosome number and habitat, specimens of Pteridiutn from southern Spain have been segregated by previous authors as a species distinct from Pteridium aquilinum , and called Pteridium herediae. However, type specimens of this taxon cannot be found. New collections of Pteridium from similar habitats in the area were examined and analysed in the present study and the results compared with those from British plants (P. aquilinum). The Spanish plants were not found to differ significantly from P. aquilinum morphologically, and all yielded chromosome counts (sporophytes, 104; gametophytes, 52) considered typical for the genus. Genotype frequency data obtained from isozyme electrophoresis of Spanish and British Pteridium yielded a genetic identity of 0.86; well within intraspecific distances found in other ferns.
These data indicate that there are currently no grounds for accepting that a form of Pteridium , distinguishable from P. aquilinum on the basis of differences in morphology, chromosome number, habitat or biochemistry, exists on the limestone soils of the Sierra de Cazorla region of southern Spain.     

The relationship between stand biomass and frond density in the clonal alga Mazzaella cornucopiae (Rhodophyta,Gigartinaceae)

The negative relationship between stand biomass and plant density observed in terrestrial plants was tested among fronds of a clonal red alga, Mazzaella cornucopiae. Stand biomass and frond density were estimated bimonthly for 1 year on 7 permanent quadrats. A positive linear correlation was found between biomass and density for the whole data set, suggesting the lack of self-thinning among fronds of this intertidal alga at natural densities. Higher frond densities could favor increased water retention among fronds, thus minimizing desiccation during low tides. In this way, stands could maintain higher production of biomass. Fronds may also be cushioned better against wave action at higher frond densities, thus decreasing the detachment of larger fronds. The temporal variation of the relationship between biomass and density was plotted separately for these 7 quadrats. Four quadrats showed positive linear correlation between both variables (the other three quadrats showed non-significant positive linear correlation). Their four slopes are statistically similar to that found for the entire data set. It is possible, then, that there is only one positive slope for the biomass-density relationship in this population. If this is true, standing biomass could be estimated from the density of fronds.     

Differential Response in the Water Status of Immature and Mature Fronds of the Ostrich Fern (Matteuccia struthiopteris [L.] Todaro) to a Mild Water Stress

Experiments were conducted in growth chambers to examine the effect of a mild water stress (−200 kilopascals polyethylene glycol) on frond elongation and water status of the ostrich fern (Matteuccia struthiopteris [L.] Todaro). Measurements were taken for two days, starting one day after the application of polyethylene glycol. Total water potential in control (well-watered) plants was always significantly higher in immature fronds than in mature fronds. The osmotic potential in mature fronds was always significantly lower (about 800 kilopascals) than in immature fronds in both control and stressed plants. In immature fronds, the stress decreased elongation and total water and pressure potentials, while in mature fronds it increased total water and pressure potentials. The decreases in total and pressure potentials in immature fronds were approximately equal to the increases in mature fronds.     

Regeneration and reproduction of Mazzaella cornucopiae (Rhodophyta, Gigartinaceae) after frond harvesting

The effects that different intensities of frond harvesting have on frond regeneration and subsequent production of reproductive structures were investigated for the red intertidal alga Mazzaella cornucopiae (Postels & Ruprecht) Hommersand from British Columbia, Canada. Harvesting was done by pruning fronds in the late spring (when stand biomass is highest) of 1993 at two intensities: total and partial collection of fronds, in this second case leaving all frond biomass less than 1 cm high in place. Holdfasts were not damaged. Total percent cover of thalli, frond density, mean frond length, and stand biomass for these experimental quadrats were statistically similar to values for control quadrats in the spring of 1994. These results suggest that one total harvest of fronds per year, done in late spring without damaging holdfasts, may give the highest sustainable yield of biomass. The effects of harvesting intensity on reproduction were variable and difficult to explain. Neither the appearance nor the abundance of cystocarpic fronds were affected by frond pruning, compared with control areas, but pruning did affect the appearance and the abundance of tetrasporic fronds. Partial pruning resulted in a longer presence of tetrasporic fronds, whereas total pruning was associated with their complete absence. Results are compared with those for the few other species of the Gigartinaceae for which experimental harvesting has been done. This revised version was published online in August 2006 with corrections to the Cover Date.     

Increased plant growth and copper uptake of host and non-host plants by metal-resistant and plant growth-promoting endophytic bacteria

The effects of inoculation with two metal-resistant and plant growth-promoting endophytic bacteria (Burkholderia sp. GL12 and Bacillus megaterium JL35) were evaluated on the plant growth and Cu uptake in their host Elsholtzia splendens and non-host Brassica napus plants grown in natural Cu-contaminated soil. The two strains showed a high level of ACC deaminase activities. In pot experiments, inoculation with strain GL12 significantly increased root and above-ground tissue dry weights of both plants, consequently increasing the total Cu uptake of E. splendens and Brassica napus by 132% and 48.2% respectively. Inoculation with strain JL35 was found to significantly increase not only the biomass of B. napus, consequently increasing the total Cu uptake of B. napus by 31.3%, but Cu concentration of E. splendens for above-ground tissues by 318% and roots by 69.7%, consequently increasing the total Cu uptake of E. splendens by 223%. The two strains could colonize the rhizosphere soils and root interiors of both plants. Notably, strain JL35 could colonize the shoot tissues and significantly increase the translocation factors and bioaccumulation factors of E. splendens. These results suggested that Burkholderia sp. GL12 and B. megaterium JL35 were valuable bacterial resource which had the potential in improving the efficiency of Cu phytoextraction by E. splendens and B. napus in a natural Cu-contaminated soil.     

Arsenic uptake and accumulation in fern species growing at arsenic-contaminated sites of southern China: field surveys

Aiming at searching for new arsenic (As) hyperaccumulators, field surveys were conducted at 12 As-contaminated sites located in Guangxi and Guangdong Provinces of southern China. Samples of 24 fern species belonging to 16 genera and 11 families as well as their associated soils were collected and As concentrations in plant and soil samples were determined by inductively coupled plasma-atomic emission spectrometry (ICP-AES). The results show that among 24 fern species, Pteris multifida and P. oshimensis can (hyper)accumulate As in their fronds with high concentrations in addition to P. vittata and P. cretica var. nervosa, which have been previously identified as As hyperaccumulators. Total As concentrations in soils associated with P. multifida and P. oshimensis varied from 1262 to 47,235 mg kg(-1), but the DTPA-extractable As concentrations were relatively low, with a maximum of 65 mg kg(-1). Forty-four of 49 samples of P. multifida collected from five sites and 3 of 13 samples of P. oshimensis collected from one site accumulated over 1000 mg As kg(-1) in their fronds and As concentrations in the fronds were higher than those in the petioles and rhizoids. Although As concentrations in the fronds of P. oshimensis (789 mg kg(-1) averaged, range 301-2142 mg kg(-1)) were comparatively lower than those of P. multifida (1977 mg kg9-1), 624-4056 mg kg(-1)), its high aboveground biomass makes it more suitable for phytoremediating As-contaminated soils. Among all the species in Pteris genus studied, Pteris semipinnata accumulated only very low As concentration in its fronds (8 mg kg(-1), 1-18 mg kg(-1)). Further research is needed to study the differences in As uptake and accumulation among fern species in the same or other genera.     

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.     

Phytoremediation potential of Pityrogramma calomelanos var. austroamericana and Pteris vittata L. grown at a highly variable arsenic contaminated site

This study examined the phytoextraction potential of two arsenic (As) hyperaccumulators, Pteris vittata L. and Pityrogramma calomelanos var. austroamericana at a historical As-contaminated cattle dip site in northern New South Wales (NSW), Australia. Total As concentration in the surface soil (0-20 cm) showed a better spatial structure than phosphate-extractable As in the surface and sub-surface soil at this site. P. calomelanos var. austroamericana produced greater frond dry biomass (mean = 130 g plant(-1)) than P. vittata (mean = 81 g plant(-1)) after 10 months of growth. Arsenic concentration and uptake in fronds were also significantly higher in P. calomelanos var. austroamericana (means = 887 mg kg(-1) and 124 mg plant(-1)) than in P. vittata (means = 674 mg kg(-1) and 57 mg plant(-1)). Our results showed that under the field conditions and highly variable soil As at the site, P. calomelanos var. austroamericana performed better than P. vittata. We predict that P. calomelanos var. austroamericana would take approximately 100 years to reduce the total As to below 20 mg kg(-1) at the site compared to > or =200 years estimated for P. vittata. However, long-term data are required to confirm these observations under field conditions.     

Effects of long-term exposure to elevated CO2 and increased nutrient supply on bracken (Pteridium aquilinum)

1. Bracken ( Pteridium aquilinum ) is an important fern with a global distribution. Little is known of the response of this species to elevated CO₂. We investigated the effects of high CO₂ (570 compared with 370 μmol mol^–1) with and without an increased nutrient supply (a combined N, P, K application) on the growth and physiology of bracken, growing in containers in controlled-environment glasshouses, over two full growing seasons. Results of growth and physiology determinations are reported for the second season.
2. Elevated CO₂ had little impact on the growth or allocation of dry mass in bracken. No significant changes were detected in dry mass of the total plant or any of the organs: rhizomes, roots and fronds. In contrast to the small effects of high CO₂, the high nutrient treatment caused a three-fold stimulation of total plant dry mass and an increase in the allocation of dry mass to above ground when compared with low nutrient controls.
3. Net photosynthetic rates in saturating light were increased by both high CO₂ and nutrient treatments, particularly in spring months (May and June). Growth in elevated CO₂ did not cause a down-regulation in light-saturated rates of photosynthesis. The increased carbon gain in the high CO₂ treatments was accompanied, in the low-nutrient plants, by higher concentrations of carbohydrates. However, in high-nutrient plants the CO₂ treatment did not cause an accumulation of carbohydrates. The absence of a growth response to elevated CO₂ in bracken despite significant increases in photosynthesis requires further investigation.     

Plant and frond dynamics of the giant kelp,Macrocystis pyrifera,forming a fringing zone in the Falkland Islands

Fluctuations in plant and frond characteristics are described for Macrocystis pyrifera (L.) C. Agardh (Laminariales, Phaeophyta) forming a fringing zone in the Falkland Islands. Giant kelp plants were sampled along a transect in the austral autumn (May 1986) and late spring (December 1986) which, according to previous frond weight analysis, were the times when extremes in population parameters were expected. Plant density and holdfast wet weights were similar for both seasons, but plants had more fronds and the fronds weighed more in spring than in autumn. Consequently, in autumn the frond biomass (1·1 wet kg m^?2) and productivity (34·1 wet g m^?2 d^?1) were lower than in spring, when a biomass of 5·0 wet kg m^?2 and a productivity of 72·4 wet g m^?2 d^?1 were recorded. Production of new fronds and loss of old fronds were determined at monthly intervals between April 1986 and March 1987. New frond production rates followed fluctuations in the quantity of light and varied between 0·08 and 0·48 fronds per plant per day. Frond loss rates did not show a seasonal pattern and fluctuated between 0·05 and 0·42 fronds per plant per day. It is suggested that the Falkland Islands Macrocystis population is more stable than most other giant kelp beds at high latitudes, because of the absence of winter storms.     

Clinal Variation of Frond Morphology and its Adaptive Implication in the Fern Ceratopteris thalictroides in Japan

Abstract Frond morphology of the highly polymorphic, annual fern Ceratopteris thalictroides was examined for local populations in south and central Japan. Among the characters examined, frond length and frond dissection were most useful for recognizing several forms. Through the comparison of these characters among populations, a geographically clinal pattern of variation was recognized; plants in higher latitudes have smaller and less dissected fronds. Plants usually grow in rice paddies during a limited period from the harvest time to the beginning of the frost season. Concordance was observed between the degree of frond length and dissection of populations and the length of the viable period in their localities. The variability of cultivated progeny of selected populations from the south and the north suggests that there are at least two ecological lines; one is genetically restricted to large and much dissected fronds and the other to smaller and less dissected fronds. The latter may be an adaptation to the short viable period in rice paddies in northern regions. High phenotypic plasticity enables these lines to adjust well the degree of frond length and dissection to the length of the viable period in each locality.