植物重金属超富集机理研究进展

植物超富集重金属机理主要涉及植物对金属离子高的吸收、运输能力,区域化作用及螯合作用等方面,其中跨膜运载蛋白的表达、调控对重金属超富集这一特性起了关键作用。金属阳离子运载蛋白家族主要包括CDF家族、NRAMP家族和ZIP家族等,在超富集植物中已克隆出多个家族的金属运载蛋白基因,这些基因的过量表达对重金属在细胞中的运输、分布和富集及提高植物的抗性方面发挥了重要作用。综述了近年来研究重金属超富集植物吸收、转运和贮存Zn、Ni、Cd等重金属的生理和分子机制所取得的主要进展。     

Lygus hesperus （Heteroptera： Miridae） tolerates high concentrations of dietary nickel

Nickel hyperaccumulator plants contain unusually elevated levels of Ni （〉 1 000 μg Ni/g）. Some insect herbivores, including Lygus hesperus （Western tarnished plant bug）, have been observed feeding on the California Ni hyperaccumulator Streptanthus polygaloides. This bug may be able to utilize S. polygaloides as a host either through its feeding behavior or by physiological tolerance of Ni. This experiment determined the Ni tolerance of L hesperus by offering insects artificial diet amended with 0, 0.4, 1, 2, 4.5, 10, 20 and 40 mmol Ni/L and recording survival. Survival varied due to Ni concentration, with diets containing 10 mmol Ni/L and greater resulting in significantly lower survival compared to the control （0 mmol Ni/L） treatment. Insects tolerated diet containing as much as 4.5 mmol Ni/L, a relatively elevated Ni concentration. I conclude that L hesperus can feed on S. polygaloides because it is Ni-tolerant, probably due to physiological mechanisms that provide it with resistance to plant chemical defenses including elemental defenses such as hyperaccumulated Ni.     

Comparative microarray analysis of Arabidopsis thaliana and Arabidopsis halleri roots identifies nicotianamine synthase, a ZIP transporter and other genes as potential metal hyperaccumulation factors

The hyperaccumulation of zinc (Zn) and cadmium (Cd) is a constitutive property of the metallophyte Arabidopsis halleri. We therefore used Arabidopsis GeneChips to identify genes more active in roots of A. halleri as compared to A. thaliana under control conditions. The two genes showing highest expression in A. halleri roots relative to A. thaliana roots out of more than 8000 genes present on the chip encode a nicotianamine (NA) synthase and a putative Zn2+ uptake system. The significantly higher activity of these and other genes involved in metal homeostasis under various growth conditions was confirmed by Northern and RT-PCR analyses. A. halleri roots also show higher NA synthase protein levels. Furthermore, we developed a capillary liquid chromatography electrospray ionization quadrupole time-of-flight mass spectrometry (CapLC-ESI-QTOF-MS)-based NA analysis procedure and consistently found higher NA levels in roots of A. halleri. Expression of a NA synthase in Zn2+-hypersensitive Schizosaccharomyces pombe cells demonstrated that formation of NA can confer Zn2+ tolerance. Taken together, these observations implicate NA in plant Zn homeostasis and NA synthase in the hyperaccumulation of Zn by A. halleri. Furthermore, the results show that comparative microarray analysis of closely related species can be a valuable tool for the elucidation of phenotypic differences between such species.     

A caterpillar (Callopistria floridensis G. (Lepidoptera: Noctuidae)) accumulates arsenic from an arsenic‐hyperaccumulating fern (Pteris vittata L.).

1. The consumption of arsenic is toxic to most biota. However, a noctuid caterpillar was recently reported feeding on a plant known to hyperaccumulate arsenic. 2. The aim of this study was to investigate the effects of arsenic‐rich Pteris vittata L. consumption by Callopistria floridensis G., and measure differences in arsenic concentrations at various stages of development (larval and adult), and associated with exuviae and frass. 3. Callopistria floridensis accumulated extraordinary concentrations of arsenic. The relative accumulation of arsenic was highest in exuviae and larvae. Larvae invariably preferred P. vittata grown on low arsenic soil to P. vittata grown on higher soil arsenic concentrations, and appeared able to selectively forage on lower arsenic concentrations within each treatment. 4. These findings show that C. floridensis is tolerant of arsenic, and successfully develops to adulthood containing elevated concentrations of arsenic. Callopistria floridensis represents the only known terrestrial animal capable of accumulating arsenic, and may have developed novel physiological and behavioural adaptations to regulate the negative effects of arsenic.     

High-nickel insects and nickel hyperaccumulator plants: A review

Insects can vary greatly in whole‐body elemental concentrations. Recent investigations of insects associated with Ni hyperaccumulator plants have identified insects with relatively elevated whole‐body Ni levels. Evaluation of the limited data available indicates that a whole‐body Ni concentration of 500 μg Ni/g is exceptional: I propose that an insect species with a mean value of 500 μg Ni/g or greater, in either larval/nymphal or adult stages, be considered a “high‐Ni insect”. Using the 500 μg Ni/g criterion, 15 species of high‐Ni insects have been identified to date from studies in Mpumalanga (South Africa), New Caledonia and California (USA). The highest mean Ni concentration reported is 3 500 μg Ni/g for nymphs of a South African Stenoscepa species (Orthoptera: Pyrgomorphidae). The majority of high‐Ni insects (66%) are heteropteran herbivores. Studies of high‐Ni insect host preference indicate they are monophagous (or nearly so) on a particular Ni hyperaccumulator plant species. Much of the Ni in bodies of these insects is in their guts (up to 66%–75%), but elevated levels have also been found in Malpighian tubules, suggesting efficient elimination as one strategy for dealing with a high‐Ni diet. Tissue levels of Ni are generally much lower than gut concentrations, but up to 1200 μg Ni/g has been reported from exuviae, suggesting that molting may be another pathway of Ni elimination. One ecological function of the high Ni concentration of these insects may be to defend them against natural enemies, but to date only one experimental test has supported this “elemental defense” hypothesis. Community‐level studies indicate that high‐Ni insects mobilize Ni into food webs but that bioaccumulation of Ni does not occur at either plant‐herbivore or herbivore‐predator steps. Unsurprisingly, Ni bioaccumulation indices are greater for high‐Ni insects compared to other insect species that feed on Ni hyperaccumulator plants. There is some evidence of Ni mobilization into food webs by insect visitors to flowers of Ni hyperaccumulator plants, but no high‐Ni insect floral visitors have been reported.     

Natural variation in cadmium tolerance and its relationship to metal hyperaccumulation for seven populations of Thlaspi caerulescens from western Europe

Thlaspi caerulescens J. & C. Presl is a distinctive metallophyte of central and western Europe that almost invariably hyperaccumulates Zn to> 1.0% of shoot dry biomass in its natural habitats, and can hyperaccumulate Ni to> 0.1% when growing on serpentine soils. Populations from the Ganges region of southern France also have a remarkable ability to accumulate Cd in their shoots to concentrations well in excess of 0.01% without apparent toxicity symptoms. Because hyperaccumulation of Cd appears to be highly variable in this species, the relationship between Cd tolerance and metal accumulation was investigated for seven contrasting populations of T. caerulescens grown under controlled conditions in solution culture. The populations varied considerably in average plant biomass (3.1‐fold), shoot : root ratio (2.2‐fold), Cd hyperaccumulation (3.5‐fold), shoot : root Cd‐concentration ratio (3.1‐fold), and shoot Cd : Zn ratio (2.6‐fold), but the degree of hyperaccumulation of Cd and Zn were strongly correlated. Two populations from the Ganges region were distinct in exhibiting high degrees of both Cd tolerance and hyperaccumulation (one requiring 3 µM Cd for optimal growth), whereas across the other five populations there was an inverse relationship between Cd tolerance and hyperaccumulation, as has been noted previously for Zn. Metal hyperaccumulation was negatively correlated with shoot : root ratio, which could account quantitatively for the differences between populations in shoot Zn (but not Cd) concentrations. On exposure to 30 µM Cd, the two Ganges populations showed marked reductions in shoot Zn and Fe concentrations, although Cd accumulation was not inhibited by elevated Zn; in the other five populations, 30 µM Cd had little or no effect on Zn or Fe accumulation but markedly reduced shoot Ca concentration. These results support a proposal that Cd is taken up predominantly via a high‐affinity uptake system for Fe in the Ganges populations, but via a lower‐affinity pathway for Ca in other populations. Total shoot Cd accumulated per plant was much more closely related to population Cd tolerance than Cd hyperaccumulation, indicating that metal tolerance may be the more important selection criterion in developing lines with greatest phytoremediation potential.     

Fast response of freshwater consumers to a new trophic resource: Predation on the recently introduced Asian bivalve Limnoperna fortunei in the lower Paraná river,South America

Abstract Comparison of Limnoperna fortunei numbers and biomass in screened (5, 15 and 40 mm) and unscreened cages deployed for 18 months in the lower Paraná delta indicates that predators harvest 26–79% (numbers), or 20–85% (biomass) of the mussel population. Predation impact decreases with mussel size. On average, 6 kg of whole live mussel × m⁻² × year⁻¹ (0.36 g of dry mussel tissue × m⁻² × day⁻¹) were eliminated from the unscreened cages. Cages with 15 and 40 mm screens lost between 1 and 2 kg × m⁻² × year⁻¹. Aquatic mammals, birds, and especially fish, are probably the main consumers of large mussels. Small L. fortunei are most probably eaten by fish and also by several invertebrates, including crustaceans, leeches and gastropods. It is suggested that L. fortunei intercepts a significant fraction of the organic carbon that the Paraná‐Uruguay rivers flush into the ocean, locally boosting numbers of benthophagous animals, deposit feeders and, indirectly, higher level predators. Our results indicate that only 15 years after its first introduction in South America this invasive species is very actively consumed by local predators, but predatory suppression of the mussel seems very unlikely. Comparisons with the effects reported for the zebra mussel (Dreissena polymorpha) in Europe and North America suggest that L. fortunei is consumed more actively and that its negative impact on the local fauna is more restricted. These differences are attributed to the fact that while D. polymorpha feeds chiefly on plankton, a limited resource, L. fortunei feeds on detrital particulate organic matter, whose supply in these large South American rivers largely exceeds consumption.     

Dynamics of Ni-based defence and organic defences in the Ni hyperaccumulator, Streptanthus polygaloides (Brassicaceae)

Plants use chemical defences to reduce damage from herbivores and the effectiveness of these defences can be altered by biotic and abiotic factors, such as herbivory and soil resource availability. Streptanthus polygaloides , a nickel (Ni) hyperaccumulator, possesses both Ni-based defences and organic defences (glucosinolates), but the extent to which these defences interact and respond to environmental conditions is unknown. S. polygaloides plants were grown on high-Ni and low-Ni soil and concentrations of Ni and glucosinolates were compared with those of the congeneric non-hyperaccumulator, S. insignus spp. insignus , grown under the same conditions. Ni contents were highest (4000 μg g⁻¹ dry tissue) in S. polygaloides plants grown on high-Ni soil. Glucosinolate content was significantly higher in S. insignus than in S. polygaloides suggesting that plants defended by Ni produce a lower concentration of organic defences. In a separate experiment, high-Ni S. polygaloides plants were exposed to simulated herbivory or live folivores to determine the inducibility of Ni-based and organic defences. Contents of Ni were not affected by either herbivory treatment, whereas glucosinolate concentrations were >30% higher in damaged plants. We concluded that the Ni-based defence of S. polygaloides is not induced by herbivory.     

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.     

Metal accumulation and its effect on leaf herbivory in an allopolyploid species Arabidopsis kamchatica inherited from a diploid hyperaccumulator A. halleri

Excessive amounts of metal ions in soil are toxic for most plant species, yet metal can also facilitate plant survival by elemental defense against herbivores and pathogens. Zinc and cadmium hyperaccumulation in Arabidopsis halleri is known to be effective for the defense against natural enemies. The allotetraploid species A. kamchatica, derived from A. halleri and a non-hyperaccumulator A. lyrata, has a lower hyperaccumulation level of zinc than A. halleri, but its significance for elemental defense remains unknown. In this study, we evaluated the accumulation levels of zinc and cadmium in the allotetraploid compared with its diploid progenitors, and evaluated the contribution of metal treatments to anti-herbivore resistance under field conditions. The accumulation level of zinc in A. kamchatica was intermediate between the progenitors, but that of cadmium was lower than in both diploid progenitors. The elemental defense of A. kamchatica and A. halleri was supported by a field experiment comparing the herbivory level between a control group and metal-supplemented plants. Moreover, the effect of elemental defense was lower in A. kamchatica than in the hyperaccumulator progenitor A. halleri, which is consistent with the metal accumulation level. This result reveals that the allotetraploid plant inherited its hyperaccumulating ability from one progenitor as an advantageous trait but at an intermediate level.     

The Arabidopsis metal tolerance protein AtMTP3 maintains metal homeostasis by mediating Zn exclusion from the shoot under Fe deficiency and Zn oversupply

Zinc ions are required to maintain the biological activity of numerous proteins. However, when mislocalized or accumulated in excess, Zn(2+) ions are toxic because of adventitious binding to proteins and displacement of other metal ions, among them Fe(2+), from their binding sites. Heterologous expression of a previously uncharacterized Arabidopsis thaliana metal tolerance protein, MTP3, in the zrc1 cot1 mutant of budding yeast restores tolerance to, and cellular accumulation of, zinc and cobalt. An MTP3-GFP fusion protein localizes to the vacuolar membrane when expressed in Arabidopsis. Ectopic over-expression of MTP3 increases Zn accumulation in both roots and rosette leaves of A. thaliana, and enhances Zn tolerance. Exposure of wild-type plants to high but non-toxic concentrations of Zn or Co, or Fe deficiency, strongly induce MTP3 expression specifically in epidermal and cortex cells of the root hair zone. Silencing of MTP3 by RNA interference causes Zn hypersensitivity and enhances Zn accumulation in above-ground organs of soil-grown plants and of seedlings exposed to excess Zn or to Fe deficiency. Our data indicate that, in wild-type A. thaliana, the AtMTP3 protein contributes to basic cellular Zn tolerance and controls Zn partitioning, particularly under conditions of high rates of Zn influx into the root symplasm.     

Nickel hyperaccumulation by Thlaspi montanum var. montanum (Brassicaceae): a constitutive trait

Adaptations to particular stresses may occur only in populations experiencing those stresses or may be widespread within a species. Nickel hyperaccumulation is viewed as an adaptation to high-Ni (serpentine) soils, but few studies have determined if hyperaccumulation ability is restricted to populations from high-Ni soils or if it is a constitutive trait found in populations on both high- and low-Ni soils. We compared mineral element concentrations of Thlaspi montanum var. montanum plants grown on normal and high-Ni greenhouse soils to address this question. Seed sources were from four populations (two serpentine, two non-serpentine) in Oregon and northern California, USA. Plants from all populations were able to hyperaccumulate Ni, showing Ni hyperaccumulation to be a constitutive trait in this species. Populations differed in their ability to extract some elements (e.g., Ca, Mg, P) from greenhouse soils. We noted a negative correlation between tissue concentrations of Ni and Zn. We suggest that the ability to hyperaccumulate Ni has adaptive value to populations growing on non- serpentine soil. This adaptive value may be a consequence of metal-based plant defense against herbivores/pathogens, metal- based interference against neighboring plant species, or an efficient nutrient scavenging system. We suggest that the Ni hyperaccumulation ability of T. montanum var. montanum may be an inadvertent consequence of an efficient nutrient (possibly Zn or Ca) uptake system.     

植物超积累重金属的生理机制研究进展

土壤重金属污染已经成为一个全球性问题。重金属超积累植物在修复土壤重金属污染中具有重要的应用前景。重金属超积累植物通常具备三个基本特征,即：根系具有从土壤中吸收重金属的强大能力、能从根到地上部分高效转运重金属、在叶片中能解毒和隔离大量重金属。本文总结了重金属超积累植物吸收、转运、隔离和解毒重金属的生理机制研究进展,以期为进一步阐明植物超积累重金属的机制及其在植物修复中的应用提供参考。