Differences in phosphorus translocation contributes to differential arsenic tolerance between plants of Borreria verticillata (Rubiaceae) from mine and non-mine sites

We have identified new arsenic-tolerant plant species Borreria verticillata (Rubiaceae) that has mine and non-mine populations at a highly contaminated site (CS) and an uncontaminated site (UCS), respectively, in Brazil. Plants of B. verticillata from both sites were cultivated at different As and P concentrations. At low P concentration, CS plants showed reduced As uptake, higher P translocation to shoots, higher constitutive levels of phenolic compounds in roots, and higher tolerance to this metalloid. At the lowest P and highest As concentration, CS plants showed higher biomass. In addition, CS plants showed higher P uptake in the absence of As, suggesting that more efficient P translocation could contribute more to tolerance than decreased As uptake. In contrast, at low P concentration, UCS plants showed higher As content in shoot and root, increase in phenol levels in roots, reduction in dry biomass, and decrease of the effective efficiency of photochemical reactions and the electron transport rate. Under higher P concentrations, the decrease in As uptake was similar in both populations. The differences between the two populations with respect to As and P uptake suggest that altered kinetic properties or expression of P transporters contribute to higher As tolerance in B. verticillata from CS. As a ruderal and As-tolerant plant, B. verticillata could be successfully used for the revegetation of contaminated soils.     

Arsenic uptake and speciation and the effects of phosphate nutrition in hydroponically grown kikuyu grass (Pennisetum clandestinum Hochst)

Background

This work focuses on the accumulation and mobility properties of arsenic (As) and the effects of phosphate (P) on its movement in Pennisetum clandestinum Hochst (kikuyu grass), grown hydroponically under increasing arsenate (As(V)) concentrations. The uptake of both ions and the relative kinetics show that phosphate is an efficient competitive inhibitor of As(V) uptake. The P/As uptake rate ratios in roots indicate that P is taken up preferentially by P/As transporters. An arsenite (As(III)) efflux from roots was also found, but this decreased when the arsenate concentration in the solution exceeded 5???M.

Methods

Increases in both arsenite and arsenate concentrations in roots were observed when the arsenate concentration in the solution was increased, and the highest accumulation of As(III) in roots was found when plants were grown at 5???M As(V). The low ratios of As accumulated in shoots compared to roots suggest limited mobility of the metalloid within Kikuyu plants.

Results

The results indicate that arsenic resistance in kikuyu grass in conditions of moderate exposure is mainly dependent on the following factors: 1) phosphate nutrition: P is an efficient competitive inhibitor of As(V) uptake because of the higher selectivity of membrane transporters with respect to phosphate rather than arsenate; and 2) a detoxification mechanism including a reduction in both arsenate and arsenite root efflux.

Conclusions

The As tolerance strategy of Kikuyu limits arsenate uptake and As translocation from roots to shoots; therefore, this plant cannot be considered a viable candidate for use in the phytoextraction of arsenic from contaminated soils or water.     

Arbuscular mycorrhiza enhanced arsenic resistance of both white clover (Trifolium repens Linn.) and ryegrass (Lolium perenne L.) plants in an arsenic-contaminated soil

In a compartmented cultivation system, white clover (Trifolium repens Linn.) and ryegrass (Lolium perenne L.), with their roots freely intermingled, or separated by 37 microm nylon mesh or plastic board, were grown together in an arsenic (As) contaminated soil. The influence of AM inoculation on plant growth, As uptake, phosphorus (P) nutrition, and plant competitions were investigated. Results showed that both plant species highly depended on mycorrhizas for surviving the As contamination. Mycorrhizal inoculation substantially improved plant P nutrition, and in contrast markedly decreased root to shoot As translocation and shoot As concentrations. It also showed that mycorrhizas affected the competition between the two co-existing plant species, preferentially benefiting the clover plants in term of nutrient acquisition and biomass production. Based on the present study, the role of AM fungi in plant adaptation to As contamination, and their potential use for ecological restoration of As contaminated soils are discussed.     

Effects of arbuscular mycorrhizal inoculation on plants growing on arsenic contaminated soil

Arbuscular mycorrhizal fungi (AMF) may play an important role in phytoremediation of As-contaminated soil. In this study the effects of AMF (Glomus mosseae, Glomus intraradices and Glomus etunicatum) on biomass production and arsenic accumulation in Pityrogramma calomelanos, Tagetes erecta and Melastoma malabathricum were investigated. Soil (243 +/- 13 microg As g(-1)) collected from Ron Phibun District, an As-contaminated area in Thailand, was used in a greenhouse experiment. The results showed different effects of AMF on phytoremediation of As-contaminated soil by different plant species. For P. calomelanos and T. erecta, AMF reduced only arsenic accumulation in plants but had no significant effect on plant growth. In contrast, AMF improved growth and arsenic accumulation in M. malabathricum. These findings show the importance of understanding different interactions between AMF and their host plants for enhancing phytoremediation of As-contaminated soils.     

Metals and arsenic in soils and corresponding vegetation at Central Elbe river floodplains (Germany)

Floodplain soils at the Elbe river are frequently polluted with metals and arsenic. High contents of these pollutants were detected down to subsoil layers. NH4NO3-extractable (phytoavailable) Cd, Ni, and Zn were elevated in horizons with high acidity. Among five common floodplain plant species, Artemisia vulgaris showed highest concentrations of Cd, Cu, and Hg, Alopecurus pratensis of As and Phalaris arundinacea of Ni, Pb, and Zn. Relationships were weak between metal concentrations in plants and phytoavailable stocks in soil. As and Hg uptake seems to be enhanced on long submerged soils. Enrichments of Cd and Hg are linked to a special plant community composition. Grassland herbage sampled in July/August revealed higher concentrations of As (+122%), Hg (+124%), and Pb (+3723%) than in May. To limit harmful transfers into the food chain, low-lying terraces and flood channels revealing highest contaminations or phytoavailabilities should be excluded from mowing and grazing.     

Assessment of macro and microelement accumulation capability of two aquatic plants

The concentrations of four macroelements (C, N, P, S) and eight trace metals (Cd, Cr, Cu, Fe, Mn, Ni, Pb, Zn) were measured in the leaves and roots of the emergent plant, Phragmites communis Trin., and in the shoots and roots of the submersed Najas marina L., taken from Lake Averno (Naples, Italy). Phragmites communis leaves showed higher concentrations of carbon, nitrogen and phosphorus than roots, while the roots exhibited significantly higher concentrations of sulphur and trace metals. Najas marina roots also showed higher concentrations of sulphur and trace metals than shoots, but these differences were less marked than in Phragmites communis except for sulphur. Sulphur was the only macronutrient to show the highest concentrations in the roots. Phragmites communis roots had higher values of Cr, Cu, Fe, Mn and Ni than Najas marina roots. By contrast, Cd, Cr, Fe, Ni, Pb and Zn concentrations were higher in Najas marina shoots than in Phragmites communis leaves. Phragmites communis, available through the year, showing high capability to accumulate trace metals in the roots, appears a good monitor of lake contamination, better than Najas marina.     

Speciation and uptake of arsenic accumulated by corn seedlings using XAS and DRC-ICP-MS

ICP-MS was used to investigate the uptake of As(III) and As(V) from hydroponics growth media by corn seedlings. It was found that arsenic uptake by the plant roots for the arsenic(V) and arsenic(III) treatments were 95 and 112 ppm, respectively. However, in the shoots of the arsenic (V) treatments had 18 ppm whereas arsenic(III) treatments had 12 ppm. XANES studies showed that As for both treatments arsenic was present as a mixture of an As(III) sulfur complex and an As(V) oxygen complex. The XANES data was corroborated by the EXAFS studies showing the presence of both oxygen and sulfur ligands coordinated to the arsenic. Iron concentrations were found to increase by 4 fold in the As(V) contaminated growth media and 7 fold in the As(III) treatment compared to the control iron concentration of 500 ppm. Whereas, the total iron concentration in the shoots was found to decrease by approximately the same amount for both treatments from 360 ppm in the control to approximately 125 ppm in both arsenic treatments. Phosphorus concentrations were found to decrease in both the roots and shoots compared to the control plants. The total sulfur in the roots was found to increase in the arsenic(III) and arsenic(V) treatments to 560 ppm and 800 ppm, respectively, compared to the control plants 358 ppm. In addition, the total sulfur in shoots of the plants was found to remain relatively constant at approximately 1080 ppm. The potassium concentrations in the plants were found to increase in the roots and decrease in the shoots.     

Fractionation and speciation of arsenic in three tea gardens soil profiles and distribution of As in different parts of tea plant (Camellia sinensis L.)

The distribution pattern and fractionation of arsenic (As) in three soil profiles from tea (Camellia sinensis L.) gardens located in Karbi-Anglong (KA), Cachar (CA) and Karimganj (KG) districts in the state of Assam, India, were investigated depth-wise (0-10, 10-30, 30-60 and 60-100 cm). DTPA-extractable As was primarily restricted to surface horizons. Arsenic speciation study showed the presence of higher As(V) concentrations in the upper horizon and its gradual decrease with the increase in soil depths, following a decrease of Eh. As fractionation by sequential extraction in all the soil profiles showed that arsenic concentrations in the three most labile fractions (i.e., water-soluble, exchangeable and carbonate-bound fractions) were generally low. Most arsenic in soils was nominally associated with the organic and Fe-Mn oxide fractions, being extractable in oxidizing or reducing conditions. DTPA-extractable As (assumed to represent plant-available As) was found to be strongly correlated to the labile pool of As (i.e. the sum of the first three fractions). The statistical comparison of means (two-sample t-test) showed the presence of significant differences between the concentrations of As(III) and As(V) for different soil locations, depths and fractions. The risk assessment code (RAC) was found to be below the pollution level for all soils. The measurement of arsenic uptake by different parts of tea plants corroborated the hypothesis that roots act as a buffer and hold back contamination from the aerial parts.     

Survival strategies of plants associated with arbuscular mycorrhizal fungi on toxic mine tailings

A field survey of metal concentrations and arbuscular mycorrhizal (AM) components of plants growing on five mining sites was conducted in Chenzhou City, Hunan Province, Southern China and a control site in Hong Kong. Significant differences were observed in the average concentrations of total heavy metals (Pb, Zn, Cu, Cd) and one metalloid (As) in contaminated soils compared with the control site. Gramineae and Compositae were the dominant plant families growing on mine tailings, with Chrysanthemum moritolium (common chrysanthemum), Cynodon dactylon (Bermuda grass), Miscanthus florodulus (Sword grass) and Pteris vittata (Ladder brake fern) commonly found at all sites. AM fungal colonization was detected in most of the plants. Comparing the four common plant species, three components of mycorrhizal colonization (arbuscules, vesicles and coiled hyphae) were found in the roots of C. dactylon and P. vittata growing at Do Shun Long (DSL) mine site. Concentrations of As in fronds were 24-fold higher than in roots of P. vittata with the highest mycorrhizal colonization rate (73%) among all sampling sites. Extensive mycorrhizal colonization (85%) was also recorded in the roots of C. dactylon with As accumulation 57 times higher than in shoots. The four common plants found in metal contaminated sites had developed different strategies for survival in the contaminated sites with the aid of indigenous AM fungi.     

Arsenic speciation, and arsenic and phosphate distribution in arsenic hyperaccumulator Pteris vittata L. and non-hyperaccumulator Pteris ensiformis L

This study examined the roles of arsenic translocation and reduction, and P distribution in arsenic detoxification of Pteris vittata L. (Chinese Brake fern), an arsenic hyperaccumulator and Pteris ensiformis L. (Slender Brake fern), a non-arsenic hyperaccumulator. After growing in 20% Hoagland solution containing 0, 133 or 267 microM of sodium arsenate for 1, 5 or 10 d, the plants were separated into fronds, rhizomes, and roots. They were analyzed for biomass, and concentrations of arsenate (AsV), arsenite (AsIII) and phosphorus. Arsenic in the fronds of P. vittata was up to 20 times greater than that of P. ensiformis, yet showing no toxicity symptoms as did in P. ensiformis. While arsenic was concentrated primarily in the fronds of P. vittata as arsenite it was mainly concentrated in the roots of P. ensiformis as arsenate. Arsenic reduction in the plants took longer than 1-d. P. vittata maintained greater P in the roots while P. ensiformis in the fronds. The high arsenic tolerance of the hyperaccumulator P. vittata may be attributed to its ability to effectively reduce arsenate to arsenite in the fronds, translocate arsenic from the roots to fronds, and maintain a greater ratio of P/As in the roots.     

Enhanced phytoremediation of arsenic contaminated land

In an attempt to clean up arsenic (As) contaminated soil, the effects of phosphorus (P) fertilizer and rhizosphere microbes on arsenic accumulation by the silverback fern, Pityrogramma calomelanos, were investigated in both greenhouse and field experiments. Field experiments were conducted in Ron Phibun District, an As-contaminated area in Thailand. Soil (136-269 microg As g(-1)) was collected there and used in the greenhouse experiment. Rhizosphere microbes (bacteria and fungi) were isolated from roots of P. calomelanos growing in Ron Phibun District. The results showed that P-fertilizer significantly increased plant biomass and As accumulation of the experimental P. calomelanos. Rhizobacteria increased significantly the biomass and As content of the test plants. Thus, P-fertilizer and rhizosphere bacteria enhanced As-phytoextraction. In contrast, rhizofungi reduced significantly As concentration in plants but increased plant biomass. Therefore, rhizosphere fungi exerted their effects on phytostabilization.     

Analysis of accumulation,extractability, and metabolization of five different phenylarsenic compounds in plants by ion chromatography with mass spectrometric detection and by atomic emission spectroscopy

Phenylated arsenic compounds occur as highly toxic contaminants in former military areas where they were formed as degradation products of chemical warfare agents. Some phenylarsenic compounds such as roxarsone and aminophenylarsonic acids were applied as food additive and veterinary drugs in stock-breeding and therefore pose an environmental risk in agricultural used sites. Very few data exist in the literature concerning uptake and effects of phenylarsenic compounds in plants growing on contaminated soils. In this study, the accumulation, extractability, and metabolization of five different phenylarsenic compounds, phenylarsonic acid, p- and o-aminophenylarsonic acid, phenylarsine oxide, and 3-nitro-4-hydroxyphenylarsonic acid called roxarsone, by the terrestrial plant Tropaeolum majus were investigated. Ion chromatography coupled to inductively coupled plasma mass spectrometry was used to differentiate these arsenic compounds, and inductively coupled plasma atomic emission spectroscopy was used for total arsenic quantification. All compounds considered were taken up by the roots and transferred to stalks, leaves, and flowers. The strongest accumulation was observed for unsubstituted phenylarsonic acid followed by its trivalent analogue phenylarsine oxide that was mostly oxidized in soil whereas the amino- or nitro- and hydroxy-substituted phenylarsonic acids were accumulated to a smaller degree.The highest extraction yield of 90% for ground leaf material was achieved by 0.1 M phosphate buffer, pH 7.7, in a two-step extraction with a total extraction time of 24 h. The extraction of higher amounts of arsenic (50–70% of total arsenic present in leaves depending on arsenic species application) from non-ground intact leaves with deionized water in comparison with the buffer (20–40% of total arsenic) is ascribed to osmotic effects. The arsenic species analysis revealed a cleavage of the amino groups from the phenyl ring for plants treated with aminophenylarsonic acids. A further important metabolic effect consisted in the production of inorganic arsenate and arsenite from the phenylated arsonic acid groups.     

Analysis of accumulation, extractability, and metabolization of five different phenylarsenic compounds in plants by ion chromatography with mass spectrometric detection and by atomic emission spectroscopy

Phenylated arsenic compounds occur as highly toxic contaminants in former military areas where they were formed as degradation products of chemical warfare agents. Some phenylarsenic compounds such as roxarsone and aminophenylarsonic acids were applied as food additive and veterinary drugs in stock-breeding and therefore pose an environmental risk in agricultural used sites. Very few data exist in the literature concerning uptake and effects of phenylarsenic compounds in plants growing on contaminated soils. In this study, the accumulation, extractability, and metabolization of five different phenylarsenic compounds, phenylarsonic acid, p- and o-aminophenylarsonic acid, phenylarsine oxide, and 3-nitro-4-hydroxyphenylarsonic acid called roxarsone, by the terrestrial plant Tropaeolum majus were investigated. Ion chromatography coupled to inductively coupled plasma mass spectrometry was used to differentiate these arsenic compounds, and inductively coupled plasma atomic emission spectroscopy was used for total arsenic quantification. All compounds considered were taken up by the roots and transferred to stalks, leaves, and flowers. The strongest accumulation was observed for unsubstituted phenylarsonic acid followed by its trivalent analogue phenylarsine oxide that was mostly oxidized in soil whereas the amino- or nitro- and hydroxy-substituted phenylarsonic acids were accumulated to a smaller degree.The highest extraction yield of 90% for ground leaf material was achieved by 0.1 M phosphate buffer, pH 7.7, in a two-step extraction with a total extraction time of 24 h. The extraction of higher amounts of arsenic (50–70% of total arsenic present in leaves depending on arsenic species application) from non-ground intact leaves with deionized water in comparison with the buffer (20–40% of total arsenic) is ascribed to osmotic effects. The arsenic species analysis revealed a cleavage of the amino groups from the phenyl ring for plants treated with aminophenylarsonic acids. A further important metabolic effect consisted in the production of inorganic arsenate and arsenite from the phenylated arsonic acid groups.     

Sequential soil washing techniques using hydrochloric acid and sodium hydroxide for remediating arsenic-contaminated soils in abandoned iron-ore mines

Sequential washing techniques using single or dual agents [sodium hydroxide (NaOH) and hydrochloric acid (HCl) solutions] were applied to arsenic-contaminated soils in an abandoned iron-ore mine area. We investigated the best remediation strategies to maximize arsenic removal efficiency for both soils and arsenic-containing washing solution through conducting a series of batch experiments. Based on the results of a sequential extraction procedure, most arsenic prevails in Fe-As precipitates or coprecipitates, and iron exists mostly in the crystalline forms of iron oxide. Soil washing by use of a single agent was not effective in remediating arsenic-contaminated soils because arsenic extractions determined by the Korean standard test (KST) methods for washed soils were not lower than 6mg kg(-1) in all experimental conditions. The results of X-ray diffraction (XRD) indicated that iron-ore fines produced mobile colloids through coagulation and flocculation in water contacting the soils, containing dissolved arsenic and fine particles of ferric arsenate-coprecipitated silicate. The first washing step using 0.2M HCl was mostly effective in increasing the cationic hydrolysis of amorphous ferrihydrite, inducing high removal of arsenic. Thus, the removal step of arsenic-containing flocs can lower arsenic extractions (KST methods) of washed soils. Among several washing trials, alternative sequential washing using 0.2M HCl followed by 1M HCl (second step) and 1M NaOH solution (third step) showed reliable and lower values of arsenic extractions (KST methods) of washed soils. This washing method can satisfy the arsenic regulation of washed soil for reuse or safe disposal application. The kinetic data of washing tests revealed that dissolved arsenic was easily readsorbed into remaining soils at a low pH. This result might have occurred due to dominant species of positively charged crystalline iron oxides characterized through the sequential extraction procedure. However, alkaline extraction using NaOH was effective in removing arsenic readsorbed onto the surface of crystalline minerals. This is because of the ligand displacement reaction of hydroxyl ions with arsenic species and high pH conditions that can prevent readsorption of arsenic.     

Arsenic accumulation and resistance mechanism in Panax notoginseng, a traditional rare medicinal herb

Panax notoginseng, a traditional rare Chinese medicinal herb, was recently found to bring health risk to consumers, mainly because soil in its major plantation area was contaminated by arsenic (As). We investigated the effect of soil As pollution on the growth and As uptake of pot-cultured P. notoginseng, and the associated mechanisms of As stressed response. Results showed that, comparing with P. notoginseng growing in a low-As soil, the root, stem, and leaf biomasses of those growing in a high-As soil significantly reduced by 0.75, 0.09 and 0.21 g seedling⁻¹, respectively. Arsenic concentrations in roots, stems and leaves of the seedlings growing in high-As soil were 22, 15 and 3 times higher than those growing in low-As soil, respectively. Regardless of the soil As concentration, As existed in plants mainly as As(III), suggesting that the reduction of As(V) is a key step in As metabolism. Arsenic was distributed primarily in cell walls (51.7% for plants growing in the low-As soil, and 51.5% in the high-As soil), followed by cytoplasm supernatant, with cell organelles containing the least As. Compared with plants growing in the low-As soil, those in the high-As soil had increased superoxide dismutase and peroxidase activities in their roots, stems, and leaves, which would be associate with improving the resistance of P. notoginseng to As stress. The results suggest that there exists some special mechanisms of As-tolerance in P. notoginseng and the study is of significance in developing measures to reduce As in the herb.     

Factors influencing arsenic accumulation by Pteris vittata: a comparative field study at two sites

This study compared the factors influencing arsenic (As) accumulation by Pteris vittata at two sites, one containing As along with Au mineralization and the other containing Hg/Tl mineralization. The soils above these two sites contained high As concentrations (26.8-2955 mg kg(-1)). Although the As concentration, pH, soil cation exchange capacity and plant biomass differed significantly between the two sites, no differences were observed in the As concentrations in the fronds and roots, or the translocation factors, of P. vittata, suggesting that this species has consistent As hyperaccumulation properties in the field. The As concentration in the fronds was positively related to phosphorus (P) and potassium (K), but negatively related to calcium (Ca), at one site. This suggested that P, K and Ca influenced As accumulation by P. vittata in the field.     

Pedological characteristics of Mn mine tailings and metal accumulation by native plants

In southern China revegetation and ecological restoration of many abandoned Mn tailings has become a major concern. To determine the major constraints for plant establishment and evaluate the feasibility of remediation, a comparative study was conducted on Mn tailings and rhizosphere soils at the boundary of the tailings pond. Both tailings and rhizosphere soils had neutral to slightly alkaline pH and normal electrical conductivity. They were both enriched with organic matter (6.8-9.2%), total N (1.77-5.94 g kg(-1)), available P (41.78-73.83 mg kg(-1)) and K (146.7-906.9 mg kg(-1)), suggesting the tailings were a nutrient rich substrate for revegetation. Mn tailings were clay textured, while rhizosphere soils were silty loam or clay loam. The compaction and anoxic nature of Mn tailings were considered to be the major constraints for plant establishment. Total Mn (31903 mg kg(-1)), Cd (119 mg kg(-1)), Cu (126 mg kg(-1)) and Zn (2490 mg kg(-1)) in tailings were all at phytotoxic levels, but did not differ significantly from those in rhizosphere soils. In both tailings and rhizosphere soils, percentages of water- and DTPA-extractable metals were less than 1% and 2% of the total metal pools, respectively. Sequential extraction revealed that the majority of Mn, Cu and Zn were associated with the residual fraction, while the majority of Cd occurred as Fe-Mn oxides. The natural succession of plants around Mn tailings formed a distinctive metal-tolerant plant community, mainly comprising nine species such as Cynodon dactylon and Humulus scandens and so on. All species studied could be good candidates for revegetation of Mn tailings.     

Uptake and accumulation behaviour of angiosperms irrigated with solutions of different arsenic species

Uptake and metabolisation of arsenic as a function of both the plant type and the chemical form of arsenic were examined. For this purpose two different plant species (Silene vulgaris and Plantago major) were selected that differed in their vitality and accumulation behaviour on arsenic-loaded substrates. The plants were cultivated on soil and irrigated with aqueous solutions of an inorganic arsenic compound (arsenious acid) and an organic compound (dimethylarsinate). The arsenic species accumulated in the parts of the plants above ground were extracted by PLE and determined using IC-ICP-MS. The concentrations and metabolisation products of arsenic found in the extracts indicate different mechanisms of arsenic uptake and transformation in both angiosperms. The arsenic species pattern showed that S. vulgaris was more arsenic--tolerable than P. major which is attributed to a low arsenate to arsenite concentration ratio in the plant compartments. S. vulgaris was also able to demethylate and reduce dimethylarsinate to form arsenite in a high extent. P. major accumulated only eight times lower concentration of arsenic, and the arsenate to arsenite concentration ratio shifted to higher values. Metabolisation products of dimethylarsinate did not occur under the present experimental conditions. The vitality of the angiosperms seems to be very dependent on the ability of the plant to reduce arsenate to arsenite.     

Unusual arsenic speciation in sea anemones

Nine species of sea anemones (Anthopleura asiatica, Actinia equina, Actinodendron arboreum, Phymanthus loligo, Entacmaea actinostoloides, Stichodactyla gigantea, S. haddoni, S. mertensii and Metridium senile) contained arsenic in the range of 1.6-7.0microg As g(-1) (wet mass basis). Irrespective of the species, water-soluble arsenic compounds accounted for more than 80% of the total arsenic. Analysis of water-soluble arsenic compounds by LC/ESI-MS revealed that four arsenicals, arsenobetaine (AB), trimethylarsoniopropionate, arsenocholine (AC) and tetramethylarsonium ion (TEMA), are contained in most species but arsenate, methylarsonic acid, dimethylarsinic acid and trimethylarsine oxide are absent in all species. Interestingly, compositional patterns of the four arsenicals greatly differed from species to species. Only three species (S. gigantea, S. haddoni and M. senile) contained AB at the highest proportions, similar to the majority of marine animals. However, the remaining six species showed unusual compositional patterns of arsenic compounds; AC was most predominant in A. arboreum and P. loligo and TEMA in A. asiatica, A. equina, E. actinostoloides and S. mertensii. On the whole, high proportions (24.6-87.1% of the water-soluble arsenic) of TEMA appear to be a peculiar characteristic of many species of sea anemones. Thus, sea anemones are an important animal group in the arsenic cycling, especially in that they may be donors of TEMA to predators.