湿地植物根表铁膜研究进展

湿地植物具有一系列生理和结构上适应浸水环境的特征,如根系的通气组织和渗氧能力,这使得植物根际微环境处于氧化状态,这样土壤中的铁和锰会在植物根表被氧化而形成红棕色的铁锰氧化物胶膜.土壤中的铁锰元素和局部的氧化环境是铁膜形成的2个重要条件.铁膜的主要组分为铁锰的氧化物及其水合物,它是一种两性胶体,可以通过离子之间的吸附-解吸、氧化-还原、有机-无机的络合等作用改变根际环境中重金属阳离子和养分的存在形态,从而影响这些离子的生物有效性,进而能影响土壤-湿地植被体系中污染物和养分的迁移.本文主要综述了湿地植物根系铁锰氧化物胶膜的形成、组成以及对植物吸收外部介质中的重金属和养分的影响,并提出了在今后的研究中仍值得进一步探讨和解决的问题.

Abstract：

Wetland plants have a series of physiological and anatomical characteristics to adapt flooding habitat, e.g. , their root aerenchyma and oxygen-penetrating capability can make the rhizospheric micro-environment be under oxidation condition, resulting in the formation of Fe/Mn plaques on root surface. Soil Fe and Mn and thee micro-environment oxidation condition are the two important prerequisites for the formation of the Fe/Mn plaques. The plaques are mainly com-posed of Fe-and Mn-0xides and their hydrates, which can alter the forms of heavy metals and nu-trients in rhizospberic micro-environment via the actions ahserption-desorption, oxidation-reduc-tion, and organic-inorganic chelation, etc. , and thereby, affect the bioavailability and the migra-tion of pollutants and nutrients in wetland soil-plant system. This paper reviewed the researches about the formation and composition of Fe/Mn plaques on wetland plant root surface, and the effects of the plaques on the heavy metals and nutrients uptake by wetland plants. Some problems worthy to be further approached were discussed.     

根表铁氧化物胶膜对水稻吸收Zn的影响

采用营养液培养方法研究了水稻根表形成的铁氧化物胶膜对水稻吸收Ｚｎ的影响．结果表明,在有Ｆｅ２＋的嫌气环境中,由于根际氧化作用水稻根表会形成红色的铁氧化物胶膜,根表的铁氧化物胶膜影响水稻对Ｚｎ的吸收．铁膜数量较少时,由于对Ｚｎ的富集作用有限,其对水稻Ｚｎ的吸收虽有促进作用,但不明显．随着根表铁膜数量的增加,这种促进作用也相应增加,并且在铁膜数量增加到一定值时,对水稻吸收Ｚｎ的促进作用达到最大．而后,随着铁膜数量的进一步增加,铁膜反而阻碍水稻对Ｚｎ的吸收,成为水稻吸收Ｚｎ的障碍层．在此过程中,水稻的根分泌物,特别是其中的植物铁载体对覆有铁膜水稻根系吸收Ｚｎ有促进作用．这种促进作用随铁膜数量的增加而逐渐减弱．因此,根表铁氧化物胶膜对水稻吸收Ｚｎ并不总是起促进作用,其作用的方向和程度取决于铁膜的数量．     

3种湿地植物对锌的吸收分配及其与根表铁氧化物胶膜的关系

在室内培养条件下,以灯心草、茭白和美人蕉3种湿地植物为材料,研究了湿地植物对锌的吸收分配能力与根表铁氧化物胶膜之间的关系.结果表明:(1)3种湿地植物积累锌的总量大小顺序为:茭白>美人蕉>灯心草,茭白积累锌的总量是灯心草的1.79倍;它们根表铁氧化物胶膜含量表现为灯心草>茭白>美人蕉,且其间存在显著差异(P<0.05).(2)锌在湿地植物中分配比例表现为;根中锌量>地上部分锌量>根表铁氧化物胶膜上吸附锌量;锌主要积累在湿地植物根中,地上部分和根表铁氧化物胶膜上吸附的锌量无显著差异.(3)湿地植物根表铁氧化物胶膜上吸附锌的数量与湿地植物地下部分锌含量呈极显著正相关(r=0.983 5**),增加根表铁氧化物胶膜上锌的数量就能明显提高地下部分锌含量;每千克土壤加入1 g FeSO4后,3种湿地植物积累锌的总量平均增加了21%.可见,湿地植物根表铁氧化物胶膜对锌的吸附也是湿地植物固定或积累锌的重要途径之一.     

植物根系对根际微环境扰动机制研究进展

根际微环境是构建植物与土壤交流沟通的桥梁,也是植物遭受胁迫时优先作出响应的区域。植物根系作为根际的主要调控者,根构型和根系分泌物种类、数量的改变均可对根际微生物和土壤动物种群分布及其结构造成影响。然而,土壤动物的扰动、微生物的分解作用也可改善根际土壤特性,提高植物抗逆性及养分利用效率,从而促进植物根系生长。可见,植物根系-根际动物-根际微生物之间存在复杂的互作关系。本文从根际内、外微环境出发,分析了根际外植物根系对微环境的物理和化学扰动、根际内植物根系与微生物的互作扰动、根际内植物根系和土壤动物的物理扰动、以植物根系分泌物为介质的化学扰动等方面研究进展,在此基础上,论述了根际微环境主要影响因子之间的互作机制,并对该领域的研究方向进行了展望。     

湿地植物根表铁膜研究进展

为了适应渍水环境,许多湿地植物都具有根系泌氧、形成铁膜的能力。因铁膜具有特殊的物理或化学结构,可以通过吸附和共沉淀作用影响元素在土壤中的化学行为和生物有效性,在植物吸收营养元素和重金属中起重要作用。综述了湿地植物根表铁膜的形成、影响因素以及根表铁膜对营养元素和重金属的生态环境效应,从表征技术方面阐述了根表铁膜的作用机制。对今后的研究方向给出如下建议:(1)扩大研究领域;(2)铁膜形成的动态变化过程;(3)铁膜对植物生理形态的影响;(4)利用先进的表征技术以确定铁膜的作用机制。     

维管植物对自然湿地甲烷排放的影响

综合评述了维管植物在自然湿地甲烷产生、氧化、传输和排放过程中的作用。维管植物光合作用的产物是甲烷产生的主要碳源,植物根系的周转和碳物质的分泌为产甲烷细菌提供底物;维管植物根际氧化是甲烷氧化最主要的途径,在植物的生长期占到总氧化量的80%左右。植物传输O2的能力和根际O2的需求是影响根际氧化的主要因素;维管植物通气组织的传输促进了甲烷从土壤向大气的输送,但所采用的传输机制影响着气体的输送效率。此外,自然湿地甲烷排放的各个过程均受到维管植物形态和植被类型的影响。维管植物在甲烷排放中的作用可以部分解释自然湿地甲烷在排放的时间(季节性变化、日变化)和空间尺度上的差异。维管植物对于自然湿地甲烷排放具有指示意义,可以用于大尺度自然湿地甲烷排放量的估算。     

湿地植物根系泌氧及其在自然基质中的扩散效应研究进展

湿地植物根系径向泌氧(ROL)是构造根际氧化-还原异质微生态系统的核心要素,其扩散层为好氧、厌氧微生物提供了良好生境并促进其代谢活动,使湿地植物根际成为有机物降解、物质循环及生命活动最为强烈的场所,已有成果证明湿地植物根系ROL的强弱与污染物的去除效果密切相关。因此,开展湿地植物根系ROL及其在自然基质中的扩散效应研究,对于了解湿地植物根系ROL机理及其根际氧环境特征,进而发挥湿地植物的污染去除功能具有十分重要的意义。基于此,首先归纳了湿地植物根系ROL特征及其受影响机制的研究现状,而后从种属差异、时空分布及对微生物的影响等方面对根系ROL在自然基质中的扩散效应国内外研究成果进行了总结,最终根据研究现状与不足对今后的研究方向进行了简要展望。创新之处在于:1)提出影响根系氧供给及氧输送释放通道的环境、生物等因素,阐述了其对根系ROL的影响机制;2)着重阐述了目前研究较少提及的根系ROL扩散效应测定方法。     

根表铁锰氧化物胶膜对不同品种水稻吸镉的影响

采用土培方法,研究了不同品种水稻吸镉的差异及其与根表铁锰氧化物胶膜的关系,结果表明：不同品种水稻其根膜,根部及地上部含镉量均存在显著性差异,且镉在不同水稻植株体内运输转移能力不同,不同水稻其根表淀积的铁锰氧化物数量也存在显著性差异,根膜及地上部的含镉量与极膜的含铁量均未达到显著性相关,但与根膜的含锰量相关性显著。     

伊贝根际微生物

<正>土壤微生物是土壤中最活跃的因子,一方面是土壤天然有机体的转化者,另一方面是土壤养分的源和库,与植物营养和土壤肥力密切相关,在土壤物质和能量循环转化过程中起着重要作用。在植物的整个生长期间,根系进行着活跃的代谢作用,向根外不断分泌有机物质,这些分泌物是根际微生物的重要营养和能量来源,其成分和数量影响着根际微生物的种类和繁殖。根际微生物的数量、活性和群落结构及其变化直接影响到植物吸收水分和养分。因此,植物、土壤和微生物之间存在着相互依赖、相互作用的复杂的三边关系[1-2]。     

根际土壤通透性对玉米水分和养分吸收的影响

为改善作物根系生长微环境,探索根际土壤通透性对作物水分和养分吸收的影响,在3个灌溉水平下（灌水量分别为每次600、400和200 ml）,采用盆栽玉米不通气、每隔2 d通1次气、每隔4 d通1次气等处理方法,研究了根际土壤通透性对盆栽玉米生理指标及水分和养分吸收的影响.结果表明：在相同灌水条件下,通气处理促进了玉米株高、叶面积的增长,提高了叶绿素含量;促进了玉米对土壤养分的吸收;显著提高了玉米的根系活力,灌水量为每次600 ml时,拔节期每隔4 d通气处理的玉米根系活力最大（8.24 mg·g^-1·h^-1）,较不通气处理（4.94 mg·g^-1·h^-1）提高了66.7%;根际土壤通气处理对盆栽玉米蒸腾的影响不显著,说明根际通气可提高玉米对水分与养分的吸收利用效率,促进植株生长发育.     

Effects of Iron and Manganese Plaques on Arsenic Uptake by Rice Seedlings (Oryza sativa L.) Grown in Solution Culture Supplied with Arsenate and Arsenite

We have shown previously that phosphorus nutrition and iron plaque on the surface of rice roots influence arsenate uptake and translocation by rice in hydroponic culture. We have now investigated the role of iron (Fe) and manganese (Mn) plaque on arsenate and arsenite uptake and translocation in rice seedlings grown hydroponically. Fe and Mn plaques were clearly visible as reddish or brown coatings on the root surface after 12 h induction, and Fe plaque was much more apparent than Mn plaque. Arsenite or arsenate supply did not decrease plant dry weights significantly. There were significant differences in shoot dry weights but little difference in root dry weights between some plaque treatments. Arsenic (As) concentrations in Fe plaque when arsenate was supplied were significantly higher than those in no plaque (control) and Mn plaque treatments, and much higher than those supplied with arsenite. This showed that Fe plaque on the rice root had higher affinity to arsenate than to arsenite. In Fe plaque treatment, the results indicated that most As was sequestered in roots when arsenite was supplied and most As concentrated in Fe plaque when arsenate was supplied. Most As was accumulated in rice roots in Mn plaque and no plaque treatments for both As species.     

Red mud (RM)-Induced enhancement of iron plaque formation reduces arsenic and metal accumulation in two wetland plant species

Human activities have resulted in arsenic (As) and heavy metals accumulation in paddy soils in China. Phytoremediation has been suggested as an effective and low-cost method to clean up contaminated soils. A combined soil-sand pot experiment was conducted to investigate the influence of red mud (RM) supply on iron plaque formation and As and heavy metal accumulation in two wetland plant species (Cyperus alternifolius Rottb., Echinodorus amazonicus Rataj), using As and heavy metals polluted paddy soil combined with three rates of RM application (0, 2%, 5%). The results showed that RM supply significantly decreased As and heavy metals accumulation in shoots of the two plants due to the decrease of As and heavy metal availability and the enhancement of the formation of iron plaque on the root surface and in the rhizosphere. Both wetland plants supplied with RM tended to have more Fe plaque, higher As and heavy metals on roots and in their rhizospheres, and were more tolerant of As and heavy metal toxicity. The results suggest that RM-induced enhancement of the formation of iron plaque on the root surface and in the rhizosphere of wetland plants may be significant for remediation of soils contaminated with As and heavy metals.     

Trace metals in mangrove seedlings: role of iron plaque formation

Metal-rich mineral deposits on the roots of aquatic plants, denominated iron plaques, may moderate the uptake of essential, but potentially toxic metals by roots. We investigated the iron plaque formation on the fine, nutritive roots of mangrove seedlings growing in contrasting environments (oxidizing sand flat sediments and reducing mangrove forest sediments) in southeast Brazil. The results indicate that Avicennia schaueriana, Laguncularia racemosa, and Rhizophora mangle seedlings developed an efficient exclusion of Fe, Mn, and Zn through iron plaque formation. This process seems to be influenced substantially by species-specific responses to environmental conditions. While Fe and Zn translocation to leaves appear to be suppressed by accumulation within root tissues, this did not appear to occur for Mn, suggesting that Mn trapping in rhizosphere sediments and iron plaque formation are the main mechanisms responsible for the Mn exclusion from the organism level. In addition to factors well recognized as affecting mangrove seedling development (e.g., salinity stress and nutrient availability), the mediation of trace metal uptake by iron plaque formation possibly contribute to determine the seedling adaptability to waterlogged conditions.     

Inoculation of Fe/Mn-oxidizing bacteria enhances Fe/Mn plaque formation and reduces Cd and As accumulation in Rice Plant tissues

Plant and Soil - Iron plaques can prevent the uptake of heavy metals on the root surface of wetland plants. The Fe and Mn oxides produced by microorganisms are major Fe/Mn-oxidizing agents in the...     

Root iron plaque formation and characteristics under N2 flushing and its effects on translocation of Zn and Cd in paddy rice seedlings (Oryza sativa)

Background and Aims

Anoxic conditions are seldom considered in root iron plaque induction of wetland plants in hydroponic experiments, but such conditions are essential for root iron plaque formation in the field. Although ferrous ion availability and root radial oxygen loss capacity are generally taken into account, neglect of anoxic conditions in root iron plaque formation might lead to an under- or over-estimate of their functional effects, such as blocking toxic metal uptake. This study hypothesized that anoxic conditions would influence root iron plaque formation characteristics and translocation of Zn and Cd by rice seedlings.

Methods

A hydroponic culture was used to grow rice seedlings and a non-disruptive approach for blocking air exchange between the atmosphere and the induction solution matrix was applied for root iron plaque formation, namely flushing the headspace of the induction solution with N₂ during root iron plaque induction. Zn and Cd were spiked into the solution after root iron plaque formation, and translocation of both metals was determined.

Key Results

Blocking air exchange between the atmosphere and the nutrient solution by N₂ flushing increased root plaque Fe content by between 11 and 77 % (average 31 %). The N₂ flushing treatment generated root iron plaques with a smoother surface than the non-N₂ flushing treatment, as observed by scanning electron microscopy, but Fe oxyhydroxides coating the rice seedling roots were amorphous. The root iron plaques sequestrated Zn and Cd and the N₂ flushing enhanced this effect by approx. 17 % for Zn and 71 % for Cd, calculated by both single and combined additions of Zn and Cd.

Conclusions

Blocking of oxygen intrusion into the nutrient solution via N₂ flushing enhanced root iron plaque formation and increased Cd and Zn sequestration in the iron plaques of rice seedlings. This study suggests that hydroponic studies that do not consider redox potential in the induction matrices might lead to an under-estimate of metal sequestration by root iron plaques of wetland plants.     

Iron sulfide formation on root surfaces controlled by the life cycle of wild rice (<Emphasis Type="Italic">Zizania palustris</Emphasis>)

Iron sulfide plaques have been observed on roots of wild rice (Zizania palustris) and other wetland plants grown in sulfur-impacted freshwater ecosystems, but the mechanism of their formation and ramifications for plants have not been investigated. We exposed a model annual wetland plant, Zizania palustris, to elevated sulfate concentrations (3.1 mM) and quantified the development of iron oxide and iron sulfide precipitates on root surfaces throughout the plant life cycle. During the onset of seed production, root surfaces amended with sulfate transitioned within 1 week from iron (hydr)oxide plaques to iron sulfide plaques. During the same week, Fe(III) decreased on roots of plants not amended with sulfate but FeS did not accumulate. Prior to FeS accumulation, sulfate-amended plants had taken up the same amount of N as unamended plants. After FeS accumulation, total plant nitrogen did not increase further on sulfate-amended plants, indicating a cessation in nitrogen uptake, whereas total plant N continued to increase in unamended plants. Sulfate-amended plants produced fewer and lighter seeds with less nitrogen than unamended plants. FeS precipitation on roots may be associated with elevated sulfide and inhibited nitrogen uptake before the end of the plant’s life cycle, thus affecting the populations of this annual aquatic plant. We propose a mechanism by which a physiologically-induced decline in radial oxygen loss near the end of a plant’s life cycle initiates a precipitous decline in redox potential at the root surface and in adjacent porewater, initiating accumulation of iron sulfide plaques. These plaques could be an important locus for iron sulfide accumulation in wetland sediments.     

Metals (Fe, Mn, Zn) in the root plaque of submerged aquatic plants collected in situ: Relations with metal concentrations in the adjacent sediments and in the root tissue

We have investigated the extent of iron oxyhydroxide deposition on the roots of two common freshwater species, Vallisneria americana Michx. and Heteranthera dubia (Jacq.) MacM., collected from different sites in the St. Lawrence River, Québec, Canada, and have related metal concentrations in the root plaques both to the geochemical conditions prevailing in the host sediments (pH; metal partitioning) and to the metal concentrations within the plant root tissue. Possible effects of root plaque on sediment geochemistry are also discussed.At those sites where the two submerged plants co-existed, the amounts of Fe deposited on their respective root surfaces were positively correlated, indicating that sediment geochemistry (pH; concentration of labile metal) exerted a more important influence on plaque formation than did inter-species differences (root physiology, morphology). Iron and Mn concentrations in the root plaque were positively correlated with each other, and with the readily extractable fractions (F1, 172) of these metals in the adjacent sediments. In contrast, Zn concentrations in the root plaque of V. americana were not related to Zn concentrations in the sediments — the dominant geochemical process at the root surface is Fe deposition, such that the quantities of Zn deposited on the roots are determined not by Zn geochemistry per se but rather by the amount of Fe deposition. Indeed the Zn/Fe ratios in the root plaque were related to the Zn/Fe ratios in the surrounding sediments (NH₂OHHCl extract).On a concentration basis (g/g), more Fe, Mn and Zn was found outside the root, in the iron plaque, than inside the root tissues. For all 3 metals, significant relationships were observed between the metal concentrations in the plaque and those inside the roots. For Zn, however, the best statistical relationship was not with [Zn]_plaque, but rather with the [Zn]/[Fe] ratio in the plaque. It is hypothesized that the Zn/Fe ratio in the root plaque reflects the free Zn²⁺ concentration adjacent to the root surface, and that this in turn affects Zn uptake by the plant root. For a given value of Zn in the sediments or in the root plaque, the Zn content of the root is inversely related to the concentration of Fe oxyhydroxides, implying that Fe plays a protective role in regulating Zn bioavailability.     

Root porosity, radial oxygen loss and iron plaque on roots of wetland plants in relation to zinc tolerance and accumulation

Background and aims

Wetland plants have been widely used in constructed wetlands for the clean-up of metal-contaminated waters. This study investigated the relationship between rate of radial oxygen loss (ROL), root porosity, Zn uptake and tolerance, Fe plaque formation in wetland plants.

Methods

A hydroponic experiment and a pot trial with Zn-contaminated soil were conducted to apply different Zn level treatments to various emergent wetland plants.

Results

Significant differences were found between plants in their root porosities, rates of ROL, Zn uptake and Zn tolerance indices in the hydroponic experiment, and concentrations of Fe and Mn on roots and in the rhizosphere in the pot trial. There were significant positive correlations between root porosities, ROL rates, Zn tolerance, Zn, Fe and Mn concentrations on roots and in the rhizosphere. Wetland plants with higher root porosities and ROL tended to have more Fe plaque, higher Zn concentrations on roots and in their rhizospheres, and were more tolerant of Zn toxicity.

Conclusions

Our results suggest that ROL and root porosity play very important roles in Fe plaque formation, Zn uptake and tolerance, and are useful criteria for selecting wetland plants for the phytoremediation of Zn-contaminated waters and soils/sediments.