The influence of arsenic chemical form and concentration on Spartina patens and Spartina alterniflora growth and tissue arsenic concentration

Arsenic (As) uptake by two perennial coastal marsh grasses growing in hydroponic conditions was studied in relation to the chemical form and concentration of As added to nutrient solution. A 4×3×2 factorial experiment was conducted with treatments consisting of four As chemical forms [arsenite, As(III); arsenate, As(V); monomethyl arsonic acid, MMAA; and dimethyl arsinic acid, DMAA], three As concentrations (0.2, 0.8, and 2.0 mg As L^-1) and two plant species (Spartina patens and Spartina alterniflora). Arsenic phytoavailability and phytotoxicity were primarily determined by the As chemical form present in the nutrient solution, though As concentration also influenced both As availability and toxicity. Application of As(V) increased root, shoot and total dry matter production; this positive plant growth response may be linked with P nutrition. Organic arsenicals and As(III) were the most phytotoxic species to both marsh grasses when plant growth was considered. Arsenic uptake and transport in plant were species-specific. Phytoavailability of As followed the trend DMAA MMAA As(V) < As(III). Root and shoot As concentrations significantly increased with increasing As application rates to the rooting medium, regardless of the As chemical form. Upon absorption, inorganic arsenicals and MMAA were mainly accumulated in the root system, while DMAA was readily translocated to the shoot.     

Transport of DMAA and MMAA into rice (Oryza sativa L.) roots

Arsenate (As(V)) transport into plant cells has been well studied. A study on rice (Oryza sativa L.) showed that arsenite is transported across the plasma membrane via glycerol transporting channels. Previous studies reported that the dimethylarsinic acid (DMAA) and monomethylarsonic acid (MMAA) uptake in duckweed (Spirodela polyrhiza L.) differed from that of As(V), and was unaffected by phosphate (H₂PO₄). This article reports the transport mechanisms of DMAA and MMAA in rice roots. Linear regression analysis showed that the DMAA and MMAA uptake in rice roots increased significantly (p ≤ 0.0002 and ≤0.0001 for DMAA and MMAA, respectively) with the increase of exposure time. Concentration-dependent influx of DMAA and MMAA showed that the uptake data were well described by Michaelis-Menten kinetics. The MMAA influx was higher than that of DMAA. The DMAA and MMAA uptake in rice roots were decreased significantly (p ≤ 0.0001 and ≤0.0077 for DMAA and MMAA, respectively) with the increase of glycerol concentration indicating that DMAA and MMAA were transported into rice roots using the same mechanisms of glycerol. Glycerol is transported into plant cells by aquaporins, and DMAA and MMAA are transported in a dose-dependent manner of glycerol which reveals that DMAA and MMAA are transported into rice roots through glycerol transporting channels. The DMAA and MMAA concentration in the solution did not affect the inhibition of their uptake rate by glycerol.     

无机砷在植物体内的吸收和代谢机制

砷污染已成为全球非常突出且急需解决的环境问题,严重威胁人类健康和环境安全.在自然环境和土壤系统中,砷的存在形态相当复杂,但植物砷毒害主要源于As（V）和As（Ⅲ）暴露.As（V）通过Pi的吸收通道被植物根系吸收,并在还原酶（AR）作用下被快速还原为As（Ⅲ）.As（Ⅲ）通过NIP蛋白通道进入植物体内,在砷甲基转移酶(ArsM)的作用下转化为甲基化砷或与谷胱甘肽（GSH）、植物螯合肽（PC）等多肽的巯基螯合封存在根部液泡或转运到地上部分,从而起到砷解毒的作用.同时,植物吸收的一部分砷也可外排到外部介质.本文以农作物尤其是水稻为主线,详述了As（V）和As（Ⅲ）吸收、外排及As（V）还原、As（Ⅲ）甲基化、螯合作用的最新研究进展,并提出了今后的研究重点.     

Genes involved in arsenic transformation and resistance associated with different levels of arsenic-contaminated soils

Background  

Arsenic is known as a toxic metalloid, which primarily exists in inorganic form [As(III) and As(V)] and can be transformed by microbial redox processes in the natural environment. As(III) is much more toxic and mobile than As(V), hence microbial arsenic redox transformation has a major impact on arsenic toxiCity and mobility which can greatly influence the human health. Our main purpose was to investigate the distribution and diversity of microbial arsenite-resistant species in three different arsenic-contaminated soils, and further study the As(III) resistance levels and related functional genes of these species.     

The response of tomato (Lycopersicon esculentum) to different concentrations of inorganic and organic compounds of arsenic

Tomato plants were cultivated in greenhouse and water solutions of arsenite (As(III)), arsenate (As(V)), methylarsonic acid (MA) and dimethylarsinic acid (DMA) were applied individually into cultivation substrate at two As levels, 5 and 15 mg kg⁻¹ of the substrate. Comparing the availability of arsenic compounds increased in order arsenite = arsenate < MA < DMA where the arsenic contents in plants decreased during vegetation period. Within a single plant, the highest arsenic concentration was found in roots followed in decreasing order by leaves, stems, and fruits regardless of arsenic compound applied. Arsenic toxicity symptoms reflected in suppressed growth of plants and a lower number and size of fruits were most significant with DMA treatment. However, the highest accumulation of arsenic by plants growing in the soil containing DMA was caused by higher mobility of this compound in the soil due to its lower sorption affinity. Our results confirmed substantial role of transformation processes of arsenic compounds in soil in uptake and accumulation of arsenic by plants.     

Methylated arsenic species in plants originate from soil microorganisms

? Inorganic arsenic (iAs) is a ubiquitous human carcinogen, and rice (Oryza sativa) is the main contributor to iAs in the diet. Methylated pentavalent As species are less toxic and are routinely found in plants; however, it is currently unknown whether plants are able to methylate As. ? Rice, tomato (Solanum lycopersicum) and red clover (Trifolium pratense) were exposed to iAs, monomethylarsonic acid (MMA(V)), or dimethylarsinic acid (DMA(V)), under axenic conditions. Rice seedlings were also grown in two soils under nonsterile flooded conditions, and rice plants exposed to arsenite or DMA(V) were grown to maturity in nonsterile hydroponic culture. Arsenic speciation in samples was determined by HPLC-ICP-MS. ? Methylated arsenicals were not found in the three plant species exposed to iAs under axenic conditions. Axenically grown rice was able to take up MMA(V) or DMA(V), and reduce MMA(V) to MMA(III) but not convert it to DMA(V). Methylated As was detected in the shoots of soil-grown rice, and in rice grain from nonsterile hydroponic culture. GeoChip analysis of microbial genes in a Bangladeshi paddy soil showed the presence of the microbial As methyltransferase gene arsM. ? Our results suggest that plants are unable to methylate iAs, and instead take up methylated As produced by microorganisms.     

Straighthead disease of rice (Oryza sativa L.) induced by arsenic toxicity

Straighthead disease is a physiological disorder of rice (Oryza sativa L.) characterized by sterility of the florates/spikelets leading to reduced grain yield. Though the exact cause of straighthead is unknown, a glass house experiment was conducted to investigate the effect of inorganic arsenic on straighthead disease in rice (Oryza sativa L.). BRRI dhan 29, a popular Bangladeshi rice strain, was grown in soils spiked with arsenic (prepared from sodium arsenate, Na₂HAsO₄·7H₂O) at the rate of 10, 20, 30, 40, 50, 60, 70, 80 and 90 mg of As kg^?1 and one control treatment was also run to compare the results. Although there may be some other soil physico-chemical factors involved, arsenic concentration was found to be closely associated with straighthead of rice. With the increase of soil arsenic concentration, the severity of straighthead increased significantly. Up to the 50 mg of As kg^?1 soil treatments, the severity of straighthead incidences were not prevalent. Straighthead resulted in sterile florets with distorted lemma and palea, reduced plant height, tillering, panicle length and grain yield. Straighthead caused approximately 17–100% sterile florates/spikelets formation and about 16–100% loss of grain yield. Straighthead also causes the reduction of panicle formation and panicle length significantly (p < 0.01). In the present study, panicle formation was found to be reduced by 21–95% by straighthead.     

Expressing ScACR3 in rice enhanced arsenite efflux and reduced arsenic accumulation in rice grains

Arsenic (As) accumulation in rice grain poses a serious health risk to populations with high rice consumption. Extrusion of arsenite [As(III)] by ScAcr3p is the major arsenic detoxification mechanism in Saccharomyces cerevisiae. However, ScAcr3p homolog is absent in higher plants, including rice. In this study, ScACR3 was introduced into rice and expressed under the control of the Cauliflower mosaic virus (CaMV) 35S promoter. In the transgenic lines, As concentrations in shoots and roots were about 30% lower than in the wild type, while the As translocation factors were similar between transgenic lines and the wild type. The roots of transgenic plants exhibited significantly higher As efflux activities than those of the wild type. Within 24 h exposure to 10 μM arsenate [As(V)], roots of ScACR3-expressing plants extruded 80% of absorbed As(V) to the external solution as As(III), while roots of the wild type extruded 50% of absorbed As(V). Additionally, by exposing the As-containing rice plants to an As-lacking solution for 24 h, about 30% of the total As derived from pre-treatment was extruded to the external solution by ScACR3-expressing plants, while about 15% of As was extruded by wild-type plants. Importantly, ScACR3 expression significantly reduced As accumulation in rice straws and grains. When grown in flooded soil irrigated with As(III)-containing water, the As concentration in husk and brown rice of the transgenic lines was reduced by 30 and 20%, respectively, compared with the wild type. This study reports a potential strategy to reduce As accumulation in the food chain by expressing heterologous genes in crops.     

Removal of selenium and arsenic by animal biopolymers

The animal biopolymers prepared from hen eggshell membrane and broiler chicken feathers, which are byproducts of the poultry-processing industry, were evaluated for the removal of the oxyanions selenium [Se(IV) and Se(VI)] and arsenic [As(III) and As(V)] from aqueous solutions. The biopolymers were found to be effective at removing Se(VI) from solution. Optimal Se(IV) and Se(VI) removal was achieved at pH 2.5–3.5. At an initial Se concentration of 100 mg/L (1.3 m M), the eggshell membrane removed approx 90% Se(VI) from the solution. Arsenic was removed less effectively than Se, but the chemical modification of biopolymer carboxyl groups dramatically enhanced the As(V) sorption capacity. Se(VI) and As(V) sorption isotherms were developed at optimal conditions and sorption equilibrium data fitted the Langmuir isotherm model. The maximum uptakes by the Langmuir model were about 37.0 mg/g and 20.7 mg/g of Se(VI) and 24.2 mg/g and 21.7 mg/g of As(V) for eggshell membrane and chicken feathers, respectively.     

Effects of two species of inorganic arsenic on the nutrient physiology of rice seedlings

This paper reports on a hydroponics experiment that was conducted to investigate the effect of inorganic arsenics on the seedlings of the rice cultivar Shanyou63. The seedlings were subjected to two treatments, i.e., As(III) and As(V). The results showed that the morphological traits of the seedlings were significantly altered after the arsenic treatments. Analysis of nitrogen, phosphorus, potassium, and arsenic contents of the roots and leaves of the seedlings indicated that the absorption of phosphorus and potassium was mainly affected by As(III), while that of nitrogen was mainly affected by As(V). The expression of 12 genes involved in the absorption and utilization of nitrogen, phosphorus, and potassium were all observed to be down-regulated after the arsenic treatments. As(V) significantly affected the absorption and utilization of nitrogen, while As(III) significantly affected those of phosphorus and potassium. The result obtained by real-time FQ-PCR regarding the difference in the gene expressions agreed with that of our hydroponics experiment.     

EXAFS study on arsenic species and transformation in arsenic hyperaccumulator

Synchrotron radiation extended X-ray absorption fine structure (SR EXAFS) was employed to study the transformation of coordination environment and the redox speciation of arsenic in a newly discovered arsenic hyperaccumulator, Cretan brake (Pteris cretica L. var nervosa Thunb). It showed that the arsenic in the plant mainly coordinated with oxygen, except that some arsenic coordinated with S as As-GSH in root. The complexation of arsenic with GSH might not be the predominant detoxification mechanism in Cretan brake. Although some arsenic in root presented as As(V) in Na₂HAsO₄ treatments, most of arsenic in plant presented as As(III)-O in both treatments, indicating that As(V) tended to be reduced to As(III) after it was taken up into the root, and arsenic was kept as As(III) when it was transported to the above-ground tissues. The reduction of As(V) primarily proceeded in the root.     

Isolation and characterization of Staphylococcus sp. strain NBRIEAG-8 from arsenic contaminated site of West Bengal

Arsenic contaminated rhizospheric soils of West Bengal, India were sampled for arsenic resistant bacteria that could transform different arsenic forms. Staphylococcus sp. NBRIEAG-8 was identified by16S rDNA ribotyping, which was capable of growing at 30,000?mg?l(-1) arsenate [As(V)] and 1,500?mg?l(-1) arsenite [As(III)]. This bacterial strain was also characterized for arsenical resistance (ars) genes which may be associated with the high-level resistance in the ecosystems of As-contaminated areas. A comparative proteome analysis was conducted with this strain treated with 1,000?mg?l(-1) As(V) to identify changes in their protein expression profiles. A 2D gel analysis showed a significant difference in the proteome of arsenic treated and untreated bacterial culture. The change in pH of cultivating growth medium, bacterial growth pattern (kinetics), and uptake of arsenic were also evaluated. After 72?h of incubation, the strain was capable of removing arsenic from the culture medium amended with arsenate and arsenite [12% from As(V) and 9% from As(III)]. The rate of biovolatilization of As(V) was 23% while As(III) was 26%, which was determined indirectly by estimating the sum of arsenic content in bacterial biomass and medium. This study demonstrates that the isolated strain, Staphylococcus sp., is capable for uptake and volatilization of arsenic by expressing ars genes and 8 new upregulated proteins which may have played an important role in reducing arsenic toxicity in bacterial cells and can be used in arsenic bioremediation.     

EXAFS study on arsenic species and transformation in arsenic hyperaccumulator

As a cost-effective, efficient and environmental friendly method for the remediation of contaminated soils and waters, phytoremediation of arsenic-con- taminated soils has drawn more and more attention[1]. The plants with the special ability to accumulate arse-nic (hyperaccumulators) are a prerequisite for phy-toremediation. Cretan brake (Pteris cretica L. var nervosa Thunb) has been shown to accumulate arsenic as much as 694 mg/kg in pinna in field investigation[2], and such elevated arsenic…     

Ecological restoration of arsenic contaminated soil by Arundo donax L

The possible arsenic tolerance mechanisms were explored in Arundo donax L. under various supplied arsenic concentrations. The treatments included control (no metal) and five doses of arsenic trioxide i.e., 0, 50, 100, 300, 600 and 1000 μg L⁻¹ As to A. donax. The phytoextraction ability of A. donax L. plants was assessed using both the translocation and bioaccumulation factors. The transpirates were collected to analyze the arsenic concentration volatilized along-with study of anatomical characteristics of the plant parts. In general, the arsenite and arsenate accumulation linearly increased in roots, shoot and leaves with the increasing supplied arsenic levels i.e., from 2.348, 2.775 and 3.25 μg g⁻¹ at 50 μg L⁻¹ to 50, 53.125 and 64.25 μg g⁻¹ arsenite, at 1000 μg L⁻¹, from 4.075, 5.425 and 13.56 μg g⁻¹ at 50 μg L⁻¹ to 71, 62.02 and 436.219 μg g⁻¹ arsenate at 1000 μg L⁻¹, respectively. The order of arsenic accumulation in A. donax L. was: solution As(III) < Root As(III) < Shoot As(III) < Leaf As(III) < Solution As(V) < Root As(V) < Shoot As(V) < Leaf As(V). The range of arsenic volatilization by A. donax L. was 7.2–22% at higher supplied arsenic (300–1000 μg L⁻¹). Volatilization was an important mechanism to avoid toxic effects of arsenic by A. donax L. in addition to bioaccumulation.     

Effects of arbuscular mycorrhiza and phosphorus application on arsenic toxicity in sunflower (Helianthus annuus L.) and on the transformation of arsenic in the rhizosphere

Effects of arbuscular mycorrhiza (AM) and phosphorus (P) application on arsenic (As) toxicity were studied in a rhizobox system with As-contaminated soil collected from Shimane Prefecture, Japan. The treatments consisted of a combination of two levels of AM (Glomus aggregatum) inoculation (−AM and +AM) and two levels of P application (−P and +P at 30 mg P kg⁻¹). Sunflower (Helianthus annuus L.) seedlings were cultured in rhizoboxes for 6 weeks. Rates of root AM infection in +AM treatments were about 40% regardless of P application. AM inoculation as well as P application reduced As toxicity symptoms, most clearly so in the +AM−P treatment. Plant growth was highest in the +AM + P treatment. Shoot As concentrations were slightly reduced by AM inoculation but enhanced by P application. Shoot P concentration in the +AM−P treatment was similar to that of +P treatments and was higher than in −AM−P. Analyses of rhizosphere soils at the end of the cultivation period indicated that P application increased water-soluble As (WS−As) in all compartments while AM inoculation increased WS−As in the central compartment only. Both the WS−arsenite [WS−As^(III)] and the dominant form, arsenate [WS−As^(V)], showed gradients toward the root surface. Dimethylarsine (DMAA) was detected in the +AM treatments only. To our knowledge, this is the first report of the occurrence of DMAA in the mycorrhizosphere. AM inoculation increased WS−P similarly as +P treatments did and promoted acid phosphatase activity in the soil. In conclusion, AM inoculation alleviated the effects of As toxicity by improving P nutrition without increasing As concentrations in the shoots. Moreover, AM appeared to be involved in the transformation of soil inorganic As into less toxic organic forms.     

Pretreatment with periodate-oxidized adenosine enhances developmental toxicity of inorganic arsenic in mice

BACKGROUND: Inorganic arsenic, given by injection to pregnant laboratory animals, can induce malformations. Arsenic methylation can be inhibited by periodate‐oxidized adenosine (PAD). Severe human health effects from high chronic arsenic exposure have mainly been reported in populations with significant levels of malnutrition, which may enhance toxicity by diminishing arsenic methylating capacity. This study sought to determine the effect of inhibition of arsenic methylation on the developmental toxicity of arsenic in a mammalian model. METHODS: PAD (100 µM/kg, i.p.), was given to pregnant CD‐1 strain mice 30min before 7.5mg/kg sodium arsenite [As(III)], i.p., or 17.9mg/kg sodium arsenate [As(V)], i.p., on gestation day 8 (GD 8; copulation plug=GD 0). Control dams received As(III), As(V), or PAD alone or were untreated. Test dams were killed on GD 17, and their litters were examined for mortality and gross and skeletal defects. RESULTS: Pretreatment with PAD before either arsenical resulted in increased maternal toxicity and lower fetal weights. Pretreatment also caused higher prenatal mortality, with 8 of 21 and 5 of 17 litters totally resorbed in the PAD plus As(III) and PAD plus As(V) treatment groups, respectively. Significant increases in the incidences of exencephaly, ablepharia, and anomalies of the vertebral centra, sternebrae, and ribs were also associated with PAD pretreatment. Short tail (3 fetuses in 3 litters) was seen only following PAD plus As(III) treatment. CONCLUSIONS: These results demonstrate that the developmental toxicity of inorganic arsenic can be enhanced by PAD, due possibly to inhibited methylation of arsenic. Birth Defects Res B 68:335–343, 2003. © 2003 Wiley‐Liss, Inc.     

The role of arsenic in obesity and diabetes

As many individuals worlwide are exposed to arsenic, it is necessary to unravel the role of arsenic in the risk of obesity and diabetes. Therefore, the present study reviewed the effects of arsenic exposure on the risk and potential etiologic mechanisms of obesity and diabetes. It has been suggested that inflammation, oxidative stress, and apoptosis contribute to the pathogenesis of arsenic-induced diabetes and obesity. Though arsenic is known to cause diabetes through different mechanisms, the role of adipose tissue in diabetes is still unclear. This review exhibited the effects of arsenic on the metabolism and signaling pathways within adipose tissue (such as sirtuin 3 [SIRT3]- forkhead box O3 [FOXO3a], mitogen-activated protein kinase [MAPK], phosphoinositide-dependant kinase-1 [PDK-1], unfolded protein response, and C/EBP homologous protein [CHOP10]). Different types of adipokines involved in arsenic-induced diabetes are yet to be elucidated. Arsenic exerts negative effects on the white adipose tissue by decreasing adipogenesis and enhancing lipolysis. Some epidemiological studies have shown that arsenic can promote obesity. Nevertheless, few studies have indicated that arsenic may induce lipodystrophy. Arsenic multifactorial effects include accelerating birth and postnatal weight gains, elevated body fat content, glucose intolerance, insulin resistance, and increased serum lipid profile. Arsenic also elevated cord blood and placental, as well as postnatal serum leptin levels. The data from human studies indicate an association between inorganic arsenic exposure and the risk of diabetes and obesity. However, the currently available evidence is insufficient to conclude that low-moderate dose arsenic is associated with diabetes or obesity development. Therefore, more investigations are needed to determine biological mechanisms linking arsenic exposure to obesity and diabetes.     

Arsenic uptake, translocation and speciation in pho1 and pho2 mutants of Arabidopsis thaliana

Arsenate [As (V)] is taken up by phosphate [P (V)] transporters in the plasma membrane of roots cells, but the translocation of As from roots to shoots is not well understood. Two mutants of Arabidopsis thaliana (L.) [( pho1 , P deficient) and ( pho2 , P accumulator)], with defects in the regulation and translocation of P (V) from roots to shoots, were therefore used in this study to investigate uptake, translocation and speciation of As in roots and shoots of plants grown in soil or nutrient solution. The shoots of the pho2 mutant contained higher P concentrations, but similar or slightly higher As concentrations, in comparison with the wild type. In the pho1 mutant, the P concentration in the shoots was lower, and the As concentration was higher, in comparison with the wild type. Both pho2 and the wild type contained mainly As (III) in roots and shoot (67–90% of total As). Arsenic was likely to be translocated by a different pathway to P (V) in the pho2 and pho1 mutants . Therefore, it is suggested that As (III) is the main As species translocated from roots to shoots in Arabidopsis thaliana.     

Effect of arsenic on visible symptom and arsenic concentration in hydroponic Japanese mustard spinach

Responses of Japanese mustard spinach (JM-spinach; Brassica rapa L. var. pervirdis) were investigated at elevated levels of arsenic (As). Plants were grown hydroponically in the greenhouse under 0, 6.7, 33.5 and 67 μM As (equal to 0, 0.5, 2.5 and 5 mg L^?1 As, respectively) for 14 days. Arsenic was used as sodium meta-arsenite (NaAsO₂). Toxicity symptom was solely shown as shoot growth repression at 33.5 and 67 μM As exposures. Dry weight (DW) enhanced by 19.4% in shoot and 38.9% in root in the 6.7 μM As level as compared to control but decreased by 48.1% and 72.1% DW in shoot and 24.1% and 61.1% DW in root in the 33.5 and 67 μM As levels, respectively. This result indicated that As at lower concentration might have slight stimulating effect on JM-spinach growth, but toxicity increased with increasing As. Based on the regression lines between growth and As concentration in the plant tissues, the critical toxicity level (CTL) of As in JM-spinach shoot was 7.85 μg g^?1 DW considering 10% DW reduction. The CTL for the root was almost 2110 μg As g^?1 DW, indicating that shoot of JM-spinach was more sensitive to As-toxicity than that of root. Arsenic concentrations increased in plant parts with increasing As in the medium. Arsenic concentrations were also compared in DW and fresh weight (FW) basis. The JM-spinach concentrated unaccepted level of As in shoots for human consumption in the higher As levels without showing visible toxicity symptom. In spite of decreasing iron (Fe) concentration in shoot in the highest As level, chlorophyll index did not decrease accordingly. Phosphorus (P) concentration also decreased. Phosphorus concentration decreased much more than Fe concentration. Low P might help to mobilize Fe in shoots, resulting in higher chlorophyll index at 67 μM As level. Phosphorus might compete with Fe in shoot tissues of As-stressed JM-spinach.     

Microbial arsenic: from geocycles to genes and enzymes

Arsenic compounds have been abundant at near toxic levels in the environment since the origin of life. In response, microbes have evolved mechanisms for arsenic resistance and enzymes that oxidize As(III) to As(V) or reduce As(V) to As(III). Formation and degradation of organoarsenicals, for example methylarsenic compounds, occur. There is a global arsenic geocycle, where microbial metabolism and mobilization (or immobilization) are important processes. Recent progress in studies of the ars operon (conferring resistance to As(III) and As(V)) in many bacterial types (and related systems in Archaea and yeast) and new understanding of arsenite oxidation and arsenate reduction by respiratory-chain-linked enzyme complexes has been substantial. The DNA sequencing and protein crystal structures have established the convergent evolution of three classes of arsenate reductases (that is classes of arsenate reductases are not of common evolutionary origin). Proposed reaction mechanisms in each case involve three cysteine thiols and S-As bond intermediates, so convergent evolution to similar mechanisms has taken place.