Influence of <Emphasis Type="Italic">Methylobacterium</Emphasis> on iron translocation in plants

Iron metabolism in plants is essential to maintain optimal growth and iron nutrition is dependent on uptake of iron from the environment and movement of iron in the plant tissues. We have examined the translocation of iron in plant leaves following foliar application of FeEDTA to Vicia faba and Zea mays. Using radiolabeled iron, we observed that iron translocation is stimulated by products of Methylobacterium mesophylicum and by the cytokinin, kinetin. When cytokinins were applied to leaves along with ⁵⁵FeEDTA, the rate of iron translocation was greater than in controls without cytokinin addition. Since recent studies indicate that M. mesophylicum is widely distributed in the environment as a pyllospheric bacterium, this organism may have an important role in enhancing translocation of nutrients in plant leaves.     

植物铁吸收、转运和调控的分子机制研究进展

铁是植物正常生命活动所必需的微量矿质元素, 铁离子的吸收、转运和利用是一个复杂的过程, 很多基因参与了这一过程。本文对近10年来发现和分离的参与植物铁吸收、转运及调控的基因研究进展进行了综述。根据最近的研究结果, 提出了植物控制铁吸收的分子调控模式(机理I)。     

植物铁吸收、转运和调控的分子机制研究进展

铁是植物正常生命活动所必需的微量矿质元素,铁离子的吸收、转运和利用是一个复杂的过程,很多基因参与了这一过程。本文对近10年来发现和分离的参与植物铁吸收、转运及调控的基因研究进展进行了综述。根据最近的研究结果,提出了植物控制铁吸收的分子调控模式(机理I)。     

The chloroplast permease PIC1 regulates plant growth and development by directing homeostasis and transport of iron

The membrane-spanning protein PIC1 (for permease in chloroplasts 1) in Arabidopsis (Arabidopsis thaliana) was previously described to mediate iron transport across the inner envelope membrane of chloroplasts. The albino phenotype of pic1 knockout mutants was reminiscent of iron-deficiency symptoms and characterized by severely impaired plastid development and plant growth. In addition, plants lacking PIC1 showed a striking increase in chloroplast ferritin clusters, which function in protection from oxidative stress by sequestering highly reactive free iron in their spherical protein shell. In contrast, PIC1-overexpressing lines (PIC1ox) in this study rather resembled ferritin loss-of-function plants. PIC1ox plants suffered from oxidative stress and leaf chlorosis, most likely originating from iron overload in chloroplasts. Later during growth, plants were characterized by reduced biomass as well as severely defective flower and seed development. As a result of PIC1 protein increase in the inner envelope membrane of plastids, flower tissue showed elevated levels of iron, while the content of other transition metals (copper, zinc, manganese) remained unchanged. Seeds, however, specifically revealed iron deficiency, suggesting that PIC1 overexpression sequestered iron in flower plastids, thereby becoming unavailable for seed iron loading. In addition, expression of genes associated with metal transport and homeostasis as well as photosynthesis was deregulated in PIC1ox plants. Thus, PIC1 function in plastid iron transport is closely linked to ferritin and plastid iron homeostasis. In consequence, PIC1 is crucial for balancing plant iron metabolism in general, thereby regulating plant growth and in particular fruit development.     

Improving iron, zinc and vitamin A nutrition through plant biotechnology

Recent understanding of plant metabolism has made it possible to increase the iron, zinc and beta-carotene (provitamin A) content in staple foods by both conventional plant breeding and genetic engineering. Improving the micronutrient composition of plant foods may become a sustainable strategy to combat deficiencies in human populations, replacing or complementing other strategies such as food fortification or nutrient supplementation.     

Iron and ferritin accumulate in separate cellular locations in <Emphasis Type="Italic">Phaseolus</Emphasis> seeds

Background  

Iron is an important micronutrient for all living organisms. Almost 25% of the world population is affected by iron deficiency, a leading cause of anemia. In plants, iron deficiency leads to chlorosis and reduced yield. Both animals and plants may suffer from iron deficiency when their diet or environment lacks bioavailable iron. A sustainable way to reduce iron malnutrition in humans is to develop staple crops with increased content of bioavailable iron. Knowledge of where and how iron accumulates in seeds of crop plants will increase the understanding of plant iron metabolism and will assist in the production of staples with increased bioavailable iron.     

Impacts of Saltwater Incursion on Plant Communities,Anaerobic Microbial Metabolism,and Resulting Relationships in a Restored Freshwater Wetland

Saltwater incursion carries high concentrations of sea salts, including sulfate, which can alter anaerobic microbial processes and plant community composition of coastal freshwater marshes. We studied these phenomena in a recently restored wetland on the coastal plain of North Carolina. We measured water inundation patterns, porewater chemistry, microbial process rates, plant tissue chemistry and iron plaque on plant roots, and quantified plant community composition across a hydrologic and salinity gradient to understand the potential interactions between saltwater incursion and changes in microbial processes and plant communities. Plant communities showed no obvious response to incursion, but were structured by inundation patterns and plant growth form (for example, graminoid versus forb). Saltwater incursion increased chloride and sulfate concentrations in surface and porewater, and drove resulting spatial patterns in anaerobic microbial metabolism rates. Plots experiencing saltwater incursion had higher sulfate reduction rates and were dominated by graminoid plant species (for example, sedges, rushes, and grasses). Graminoid plant species’ roots had greater iron plaque formation than forb and submerged species, indicative that graminoid plant species are supplying more oxygen to the rhizosphere, potentially influencing microbial metabolism. Future studies should focus on how plant and microbial communities may respond to saltwater incursion at different time scales, and on parsing out the influence that plants and microbes have on each other as freshwater wetlands experience sea level rise.     

Organic acids and Fe deficiency: a review

Organic acid concentrations often increase with iron deficiency in different plant parts such as roots, leaves and stem exudates. The review summarises data available on the changes in the concentrations of organic anions in plants with iron deficiency and the effects of these changes in plant metabolism. The paper reviews data available in the literature on the changes in xylem and apoplasmic fluid composition with iron deficiency, both in plants grown in controlled conditions and in the field, and discusses the possible ways of iron complexation and transport in these compartments. The characteristics of the iron reduction and uptake by the iron-deficient leaf mesophyll cells are also discussed, with especial emphasis in the possible roles of organic acids in these processes. Both the possible causes and functions of the organic acid concentration increases in iron-deficient plants are reviewed.     

有机酸代谢在植物适应养分和铝毒胁迫中的作用

有机酸不仅是碳代谢的中间体,而且在一些植物应对养分缺乏、金属胁迫以及根-土界面中操纵植物-微生物间的交互作用方面都发挥着关键作用.从植物营养学角度,对最近关于植物体内有机酸的形成与生理,及其与氮素代谢,磷和铁的吸收,铝的耐受以及土壤生态之间的关系等方面进行了总结,并对有机酸的跨膜运输、转基因模型中有机酸调控的生物技术操作的最新发现进行了讨论,以期为理解有机酸代谢的植物营养学基本原理提供基础.最后,还提出了通过生物技术,培育作物新品种,以更好地适应环境与金属胁迫.     

Intra- and intercellular iron trafficking and subcellular compartmentation within roots

Iron is abundant in most soils, but ferric compounds are almost insoluble. Therefore, plant roots use as tools acidification and enzymatic reduction of iron at the outer cell surface (strategy I) or solubilization by phytosiderophores, which are specific ferric chelators (strategy II). In the first case, iron is taken up as Fe²⁺ into the root symplast, and in the latter one, iron is taken up as Fe(III) complex. The path of iron from the root surface, up to the point of the xylem vessels within the central cylinder, may be completely symplasmic. However, a part of this route also may be an apoplasmic one, through the free space of the cell walls of rhizodermis and cortex (apoplast). In the endodermis, the Casparian band forms a strict barrier for apoplasmic transport; to move past this site, all ions must enter the symplast. During symplasmic transport, the intracellular environment is protected against the reactive species of iron by handling of iron in chelated forms. A promising candidate for this purpose is the plant-endogenous chelator nicotianamine. At the apoplasmic site, iron can be oxidized followed by precipitation as hydroxide or phosphate compounds. Thus, a pool of apoplastic iron can be formed, as shown by reductive mobilization or by proton-induced X-ray emission. This pool may be remobilized when iron deficiency takes place. During radial transport to the vessels, vacuoles may compete with the transport stream forming an iron store. When there is an iron excess, as in plants growing in waterlogged soils or by experimental techniques, plants can escape the deleterious effects of free iron by depositing it in phytoferritin, a storage protein inducible under iron excess. Also, nicotianamine forms a pool of metabolically available iron. Thus, in roots cells of the nicotianamine-free tomato mutant chloronerva iron precipitations occur as evidenced by energy dispersive X-ray analysis and the electron microscopic energy loss technique of energy spectroscopic imaging. Future research concerning the plant root's iron metabolism are needed to clarify the function of nicotianamine in intra- and intercellular iron trafficking and to identify the so-called iron-sensor which mediates the regulation of iron acquisition reactions of rhizodermal cells in response to the iron nutritional status of the plant.     

Maize ZmFDR3 localized in chloroplasts is involved in iron transport

Iron is an essential nutrient for plant metabolism such that Fe-limited plants display chlorosis and suffer from reduced photosynthetic efficiency. Differential display previously identified genes whose expression was elevated in Fe-deficient maize roots. Here,we describe the functional characterization of one of the genes identified in the screen,ZmFDR3 (Zea maize Fe-deficiency-related). Heterologous functional complementation assays using a yeast iron uptake mutant showed that ZmFDR3 functions in iron tra...     

Cellular regulation of iron assimilation

Cells of plants, most microorganisms, and animals require well-defined amounts of iron for survival, replication, and differentiation. The metal is an important component of such processes as synthesis of DNA, RNA, and chlorophyll; electron transport; oxygen metabolism; and nitrogen fixation. Because of the insolubility of iron in aerobic environments at neutral and alkaline pH values, cells have had to devise specific strategies to assimilate the metal. These include (1) development of systems for reducing ferric ions to the more soluble ferrous ions at the cell surface, (2) employment of small carrier molecules (termed siderophores) that have high affinity for ferric ions and receptor proteins for the ferrated molecules, and (3) use of transferrin and other proteins that can transport ferric ions. Excessive amounts of iron are toxic, however, and intracellular storage capacity is limited and efflux mechanisms generally are lacking. Thus, cells have had to develop methods of preventing over-accumulation of the metal. These include use of (1) oxygen to convert ferrous to ferric ions, (2) small molecules that can bind ferrous ions, termed siderophraxes, and (3) proteins that, when combined with ferrous ions, repress the expression of iron transport genes. Often, one organism can prevent growth of neighbors by restricting their access to iron. In other cases, cells assist each other by sharing iron acquisition systems or by restricting influx of excess iron. Homeostatic control of other essential trace metals also is required for optimal cell function. Nevertheless, since iron thus far has received most attention, it serves as the model of mineral metabolism. Moreover, many of the observations made on control of iron metabolism suggest possible applications in prevention and management of plant and animal infections as well as of neoplastic diseases, arthropathy, and cardiomyopathy. This review will focus on (1) problems at the cellular level of iron acquisition, storage, and exclusion; and (2) the strategies devised by cells of plants, microorganisms, and animals to solve these problems.     

Current understanding of the regulation of methionine biosynthesis in plants

Plants can provide most of the nutrients for the human diet. However, the major crops are often deficient in some of the nutrients. Thus, malnutrition, with respect to micronutrients such as vitamin A, iron, and zinc, but also macronutrients such as the essential amino acids lysine and methionine, affects more than 40% of the world's population. Recent advances in molecular biology, but also the grasp of biochemical pathways, metabolic fluxes, and networks can now be exploited to produce crops enhanced in key nutrients to increase the nutritional value of plant-derived foods and feeds. Some of the predictions appear to be accurate, while others not, reflecting the fact that plant metabolism is more complex than presently understood. A good example for a complex regulation is the methionine biosynthetic pathway in plants. The nutritional importance of Met and cysteine has motivated extensive studies of their roles in plant molecular physiology, especially regarding to their transport, synthesis, and accumulation in plants. Recent studies have demonstrated that Met metabolism is regulated differently in various plant species.     

Phosphate deficiency promotes modification of iron distribution in Arabidopsis plants

Phosphate (Pi) is an essential element for plant development and metabolism. Due to its low availability and mobility in soils, it is often a limiting nutrient for their growth. This phenomenon is reinforced by the formation of insoluble complexes in the environment with many cations, affecting the solubility of both phosphate and associated ions. This interaction is investigated here for iron, a strong phosphate chelator. Depleting the medium in phosphate clearly resulted in an increase of iron content in Arabidopsis. These modifications triggered molecular responses linked with iron status (transport, homeostasis and accumulation). Interestingly, physiological modifications affecting iron storage were also observed. The accumulation of phosphate/iron complexes in the vacuoles of plants grown in Pi-rich medium disappeared in Pi-depleted medium in favor of accumulation of iron inside the chloroplasts, likely associated with ferritin.     

Zinc toxicity in corn as a result of a geochemical anomaly

Summary Corn grown on zinc-rich soil (adjacent to an abandoned zinc mine) showed severe chlorosis and stunting. Soil zinc content was positively correlated with leaf zinc content, but not correlated with leaf iron content. Soil zinc was negatively correlated, and soil iron positively correlated, with chlorophyll content. Excess zinc may interfere with iron metabolism in the plant, but does not appear to affect the iron supply to the leaf.     

Nicotianamine: mediator of transport of iron and heavy metals in the phloem?

Recent work has demonstrated that minerals in plants are circulated between root and shoot. This occurs during the whole life time and renders possible response to changing environmental conditions. This mineral circulation occurs through intensive solute exchange between xylem and phloem in roots, stems, and leaves. The transport form of heavy metals such as iron, manganes, zinc and copper in the phloem, whether ionic or chelated, is unclear in most cases.
The unusual amino acid nicotianamine (NA) is ubiquitous throughout the plant kingdom. It is a chelator of several divalent transition metals. Its physiological role was investigated with the tomato mutant chloronerva, the only known NA-free multicellular plant. The mutant also exhibits disturbances of its iron metabolism and that of other heavy metals. This leads, among others, to a typical intercostal chlorosis and progressive iron accumulation in the leaves. From the heavy metal chelating properties of NA and from the phenotype of the mutant chloronerva it is concluded that NA is needed for normal distribution of heavy metals in young growing tissues fed via the phloem. This function could be fulfilled by mediating phloem loading or unloading of heavy metals as well as by preventing their precipitation in the alkaline phloem sap. An attempt is made to explain the chloronerva phenotype in the light of the phloem transport hypothesis of chelated iron.     

Dissecting plant iron homeostasis under short and long-term iron fluctuations

A wealth of information on the different aspects of iron homeostasis in plants has been obtained during the last decade. However, there is no clear road-map integrating the relationships between the various components. The principal aim of the current review is to fill this gap. In this context we discuss the lack of low affinity iron uptake mechanisms in plants, the utilization of a different uptake mechanism by graminaceous plants compared to the others, as well as the roles of riboflavin, ferritin isoforms, nitric oxide, nitrosylation, heme, aconitase, and vacuolar pH. Cross-homeostasis between elements is also considered, with a specific emphasis on the relationship between iron homeostasis and phosphorus and copper deficiencies. As the environment is a crucial parameter for modulating plant responses, we also highlight how diurnal fluctuations govern iron metabolism. Evolutionary aspects of iron homeostasis have so far attracted little attention. Looking into the past can inform us on how long-term oxygen and iron-availability fluctuations have influenced the evolution of iron uptake mechanisms. Finally, we evaluate to what extent this homeostastic road map can be used for the development of novel biofortification strategies in order to alleviate iron deficiency in human.