Rhizosphere geometry and heterogeneity arising from root-mediated physical and chemical processes

The rhizosphere differs from the bulk soil in a range of biochemical, chemical and physical processes that occur as a consequence of root growth, water and nutrient uptake, respiration and rhizodeposition. These processes also affect microbial ecology and plant physiology to a considerable extent. This review concentrates on two features of this unique environment: rhizosphere geometry and heterogeneity in both space and time. Although it is often depicted as a soil cylinder of a given radius around the root, drawing a boundary between the rhizosphere and bulk soil is an impossible task because rhizosphere processes result in gradients of different sizes. For instance, because of diffusional constraints, root uptake can result in a depletion zone extending <1 mm for phosphate to several centimetres for nitrate, while respiration may affect the bulk of the soil. Rhizosphere processes are responsible for spatial and temporal heterogeneities in the soil, although these are sometimes difficult to distinguish from intrinsic soil heterogeneity. A further complexity is that these processes are regulated by plants, microbial communities and soil constituents, and their many interactions. Novel in situ techniques and modelling will help in providing a holistic view of rhizosphere functioning, which is a prerequisite for its management and manipulation.     

Soil factors controlling mineral N uptake by Picea engelmannii seedlings: the importance of gross NH4+ production rates

* Hydroponic studies suggest that plant nitrogen (N) demand determines the rate of mineral N uptake; however, field observations show N limitation to be widespread. Field experiments are needed to understand soil factors controlling mineral N uptake. * We planted Picea engelmannii seedlings that had initially been grown from sterilized seeds, on a recently clearcut site. We applied a hybrid isotope dilution/pulse labelling technique to compare the gross production rate, concomitantly to the plant uptake rate, of soil mineral N. We also measured mineral N concentrations, microbial N, and percent ectomycorrhizal root tips. * Gross NH4+ production rate was the most important determinant of plant uptake rate. Exploratory path analysis suggested that plant uptake was also determined by microbial N, which was, in turn, determined by soil mineral N concentrations. Percent ectomycorrhizal root tips was negatively related to gross NO3- production rate and microbial N concentrations. * We conclude that nutrient flux density is important in controlling plant uptake. Mycorrhizal colonization may alter N dynamics in the rhizosphere without affecting mineral N uptake by seedlings.     

The role of fungi in biogenic weathering in boreal forest soils

In this article we discuss the possible significance of biological processes, and of fungi in particular, in weathering of minerals. We consider biological activity to be a significant driver of mineral weathering in forest ecosystems. In these environments fungi play key roles in organic matter decomposition, uptake, transfer and cycling of organic and inorganic nutrients, biogenic mineral formation, as well as transformation and accumulation of metals. The ability of lichens, mutualistic symbioses between fungi and photobionts such as algae or cyanobacteria, to weather minerals is well documented. The role of mycorrhizal fungi forming symbioses with forest trees is less well understood, but the mineral horizons of boreal forests are intensively colonised by mycorrhizal mycelia which transfer protons and organic metabolites derived from plant photosynthates to mineral surfaces, resulting in mineral dissolution and mobilisation and redistribution of anionic nutrients and metal cations. The mycorrhizal mycelia, in turn provide efficient systems for the uptake and direct transport of mobilised essential nutrients to their host plants which are large sinks. Since almost all (99.99 %) non-suberised lateral plant roots involved in nutrient uptake are covered by ectomycorrhizal fungi, most of this exchange of metabolites must take place through the plant–fungus interface. This idea is still consistent with a linear relationship between soil mineral surface area and weathering rate since the mycelia that emanate from the tree roots will have a larger area of contact with minerals if the mineral surface area is higher. Although empirical models based on bulk soil solution chemistry may fit field data, we argue that biological processes make an important contribution to mineral weathering and that a more detailed mechanistic understanding of these must be developed in order to predict responses to environmental changes and anthropogenic impact.     

施用污泥对油菜根际养分和不同种类重金属的影响

根际是控制植物养分动态的重要因素,养分动态也影响着根际土壤环境。当土壤被污水污泥改良后,根际土壤中的养分和重金属性质也会发生变化。目前很少有人研究施用污泥的土壤中植物根系对根际重金属有效性和分布的影响。采用根垫—冰冻薄层切片法对施用污泥后土壤中油菜根际的养分和重金属分布情况进行研究,以期探明污泥改良土壤中根际重金属的活化特征。当土壤施用污泥后,根际土壤中DTPA提取态Zn,Cd,Ni,Mn,有效磷,有效钾和铵态氮被显著消耗,而根际土壤中DTPA提取态Cu没有明显的消耗或积累。当土壤中施用大量污泥时,根际土壤的pH值随着离根表面距离的增加而增加。无论土壤是否用污泥处理,油菜根际土壤中可交换态Cu都显著减少。当土壤被50%污泥改良时,在距离根表面0—2 mm处的油菜根际土壤中碳酸盐结合态,铁锰氧化物结合态,有机物结合态,残渣态的Cu和Zn都被消耗较多。污泥的施用对油菜的生长有促进作用。随着污泥施用量的增加,油菜地上部分Cu和Zn的含量没有显著变化。施用污泥量小于25%的土壤中,污泥没有增加重金属的可利用性和移动性。除了Cu,油菜根际土壤中DTPA提取态Zn,Cd,Ni的减少表明施用污泥的土壤中重金属的活化是非常有限的。     

Root exudates as mediators of mineral acquisition in low-nutrient environments

Plant developmental processes are controlled by internal signals that depend on the adequate supply of mineral nutrients by soil to roots. Thus, the availability of nutrient elements can be a major constraint to plant growth in many environments of the world, especially the tropics where soils are extremely low in nutrients. Plants take up most mineral nutrients through the rhizosphere where micro-organisms interact with plant products in root exudates. Plant root exudates consist of a complex mixture of organic acid anions, phytosiderophores, sugars, vitamins, amino acids, purines, nucleosides, inorganic ions (e.g. HCO₃ ^–, OH^–, H⁺), gaseous molecules (CO₂, H₂), enzymes and root border cells which have major direct or indirect effects on the acquisition of mineral nutrients required for plant growth. Phenolics and aldonic acids exuded directly by roots of N₂-fixing legumes serve as major signals to Rhizobiaceae bacteria which form root nodules where N₂ is reduced to ammonia. Some of the same compounds affect development of mycorrhizal fungi that are crucial for phosphate uptake. Plants growing in low-nutrient environments also employ root exudates in ways other than as symbiotic signals to soil microbes involved in nutrient procurement. Extracellular enzymes release P from organic compounds, and several types of molecules increase iron availability through chelation. Organic acids from root exudates can solubilize unavailable soil Ca, Fe and Al phosphates. Plants growing on nitrate generally maintain electronic neutrality by releasing an excess of anions, including hydroxyl ions. Legumes, which can grow well without nitrate through the benefits of N₂ reduction in the root nodules, must release a net excess of protons. These protons can markedly lower rhizosphere pH and decrease the availability of some mineral nutrients as well as the effective functioning of some soil bacteria, such as the rhizobial bacteria themselves. Thus, environments which are naturally very acidic can pose a challenge to nutrient acquisition by plant roots, and threaten the survival of many beneficial microbes including the roots themselves. A few plants such as Rooibos tea (Aspalathus linearis L.) actively modify their rhizosphere pH by extruding OH^– and HCO₃ ^– to facilitate growth in low pH soils (pH 3 – 5). Our current understanding of how plants use root exudates to modify rhizosphere pH and the potential benefits associated with such processes are assessed in this review.     

Changes in the extractability of cations (Ca,Mg and K) in the rhizosphere soil of Norway spruce (Picea abies) roots

Nutrient concentrations in the rhizosphere soil can be higher, lower or remain unchanged compared to the bulk soil, but relatively little is known about such changes for basic cations in the rhizosphere of tree roots. A modified root container technique of studying rhizosphere processes was employed. Plexiglas cylinders were horizontally split by a membrane with 30 M mesh size into an upper compartment for root growth and a root-free lower compartment, each with an inner diameter of 5 cm and a height of 10 cm. One 2-year-old Norway spruce (Picea abies) seedling was transplanted from a nursery into each cylinder. Plants were not specifically inoculated, but roots were colonised by a mix of ectomycorrhizal fungi originating from the nursery. The nutrient poor mineral soil used in the experiment was taken from a forest site in Bayerischer Wald, southern Germany. The soil was either supplied with a mix of Ca, Mg and K, or not supplied with these cations. Plants were harvested 30 weeks after transplanting. The nylon membrane between the root compartments restricted root growth to the upper compartment, so that by the end of the experiment a root mat was formed at the top side of the membrane. In the lower compartment, soil nearest to the root mat was regarded as rhizosphere soil while soil in a distance from the root mat was regarded as bulk soil. In the upper compartment, rhizosphere soil was obtained at the end of the experiment by gently shaking the roots. The soils were analysed for Ca, Mg and K contents following two different soil extraction methods. In the fertilised treatment, H₂O-extractable Ca and Mg were accumulated in the rhizosphere. In contrast, K (NH₄Cl-extraction) was depleted in the rhizosphere. In the bottom tube, the depletion of K (NH₄Cl-extraction) was restricted to 1 cm distance from the root mat. In unfertilised soil, Ca, Mg and K concentrations did not differ clearly between rhizosphere and bulk soils. The results indicated that the occurrence of cation gradients in the rhizosphere depended on the level of soil nutrient supply. Distinct rhizosphere effects were measured by conventional soil extraction methods only when the soil was freshly fertilised with mineral elements prior to the experiment. In this case, K depletion in the rhizosphere reflected higher K uptake by the fertilised Norway spruce plants. For low-nutrient soils, novel techniques are required to follow subtle changes in the rhizosphere.     

Measured and modelled differences in nutrient concentrations between rhizosphere and bulk soil in a Norway spruce stand

The gradient in soil characteristics from the bulk soil to the root surface is important to roots and to the organisms that live in the rhizosphere. Our ability to measure ion concentrations at the root surface is extremely limited, and models are largely untested. We used data from a well studied Norway spruce stand in SW Sweden to compare the measured difference in nutrient concentrations between rhizosphere and bulk soil with the difference predicted by a steady-state simulation model based on ecosystem budgets of nutrient uptake. The simulation model predicted depletion of NH₄, Ca, Mg, K in the rhizosphere, which shows that budgeted uptake rates were greater than the mass flow of bulk solution towards the root. In plots treated with ammonium sulphate, the model predicted an accumulation of S in the rhizosphere. In contrast, the observed rhizosphere concentrations were generally enriched in nutrients, relative to bulk soil. Collecting rhizosphere soil adhering to root surfaces may not be an appropriate method for describing the concentration gradient around the root. In addition, the simulation model omits some processes affecting conditions in the rhizosphere that are important to explaining nutrient uptake.     

Three-dimensional visualization and quantification of water content in the rhizosphere

? Despite the importance of rhizosphere properties for water flow from soil to roots, there is limited quantitative information on the distribution of water in the rhizosphere of plants. ? Here, we used neutron tomography to quantify and visualize the water content in the rhizosphere of the plant species chickpea (Cicer arietinum), white lupin (Lupinus albus), and maize (Zea mays) 12 d after planting. ? We clearly observed increasing soil water contents (θ) towards the root surface for all three plant species, as opposed to the usual assumption of decreasing water content. This was true for tap roots and lateral roots of both upper and lower parts of the root system. Furthermore, water gradients around the lower part of the roots were smaller and extended further into bulk soil compared with the upper part, where the gradients in water content were steeper. ? Incorporating the hydraulic conductivity and water retention parameters of the rhizosphere into our model, we could simulate the gradual changes of θ towards the root surface, in agreement with the observations. The modelling result suggests that roots in their rhizosphere may modify the hydraulic properties of soil in a way that improves uptake under dry conditions.     

Nutrient composition of Douglas-fir rhizosphere and bulk soil solutions

Rhizosphere soil solution is the direct source of nutrients for plant uptake. The nutrient composition of rhizosphere soil solution can be very different from that of bulk soil solution due to root exudation, nutrient uptake and rhizosphere microorganism activity. This study examined the nutrient composition of Douglas-fir rhizosphere soil solution in two soils belonging to the Nisqually and Pitcher soil series and compared rhizosphere solution with that of bulk soil solution. Fertilized and unfertilized Nisqually soils were also compared. Soil solutions were collected using centrifugation. Results indicated that nutrient concentrations in the rhizosphere solutions were typically higher than that of bulk soil solutions when no fertilizer was applied. Differences in the concentrations of nutrients between the rhizosphere and bulk soil solutions were masked by the addition of fertilizers. Rhizosphere solution pH also appeared to be affected by the concentration of NH₄ and NO₃ in the solution. With a higher concentration of NH₄ relative to NO₃ in the rhizosphere soil solution, the solution pH of the rhizosphere was lower than that of the bulk soil, but with a lower concentration of NH₄ relative to NO₃, the solution pH of the rhizosphere was higher than that of the bulk soil solution.     

Origins of root-mediated pH changes in the rhizosphere and their responses to environmental constraints: A review

The aim of the present review is to define the various origins of root-mediated changes of pH in the rhizosphere, i.e., the volume of soil around roots that is influenced by root activities. Root-mediated pH changes are of major relevance in an ecological perspective as soil pH is a critical parameter that influences the bioavailability of many nutrients and toxic elements and the physiology of the roots and rhizosphere microorganisms. A major process that contributes root-induced pH changes in the rhizosphere is the release of charges carried by H⁺ or OH^– to compensate for an unbalanced cation–anion uptake at the soil–root interface. In addition to the ions taken up by the plant, all the ions crossing the plasma membrane of root cells (e.g., organic anions exuded by plant roots) should be taken into account, since they all need to be balanced by an exchange of charges, i.e., by a release of either H⁺ or OH^–. Although poorly documented, root exudation and respiration can contribute some proportion of rhizosphere pH decrease as a result of a build-up of the CO₂ concentration. This will form carbonic acid in the rhizosphere that may dissociate in neutral to alkaline soils, and result in some pH decrease. Ultimately, plant roots and associated microorganisms can also alter rhizosphere pH via redox-coupled reactions. These various processes involved in root-mediated pH changes in the rhizosphere also depend on environmental constraints, especially nutritional constraints to which plants can respond. This is briefly addressed, with a special emphasis on the response of plant roots to deficiencies of P and Fe and to Al toxicity. Finally, soil pH itself and pH buffering capacity also have a dramatic influence on root-mediated pH changes.     

Effects of plants and microorganisms in constructed wetlands for wastewater treatment

Constructed wetlands are a natural alternative to technical methods of wastewater treatment. However, our understanding of the complex processes caused by the plants, microorganisms, soil matrix and substances in the wastewater, and how they all interact with each other, is still rather incomplete. In this article, a closer look will be taken at the mechanisms of both plants in constructed wetlands and the microorganisms in the root zone which come into play when they remove contaminants from wastewater. The supply of oxygen plays a crucial role in the activity and type of metabolism performed by microorganisms in the root zone. Plants' involvement in the input of oxygen into the root zone, in the uptake of nutrients and in the direct degradation of pollutants as well as the role of microorganisms are all examined in more detail. The ways in which these processes act to treat wastewater are dealt with in the following order: Technological aspects; The effect of root growth on the soil matrix; Gas transport in helophytes and the release of oxygen into the rhizosphere; The uptake of inorganic compounds by plants; The uptake of organic pollutants by plants and their metabolism; The release of carbon compounds by plants; Factors affecting the elimination of pathogenic germs.     

Carbon flow in the rhizosphere: carbon trading at the soil–root interface

The loss of organic and inorganic carbon from roots into soil underpins nearly all the major changes that occur in the rhizosphere. In this review we explore the mechanistic basis of organic carbon and nitrogen flow in the rhizosphere. It is clear that C and N flow in the rhizosphere is extremely complex, being highly plant and environment dependent and varying both spatially and temporally along the root. Consequently, the amount and type of rhizodeposits (e.g. exudates, border cells, mucilage) remains highly context specific. This has severely limited our capacity to quantify and model the amount of rhizodeposition in ecosystem processes such as C sequestration and nutrient acquisition. It is now evident that C and N flow at the soil–root interface is bidirectional with C and N being lost from roots and taken up from the soil simultaneously. Here we present four alternative hypotheses to explain why high and low molecular weight organic compounds are actively cycled in the rhizosphere. These include: (1) indirect, fortuitous root exudate recapture as part of the root’s C and N distribution network, (2) direct re-uptake to enhance the plant’s C efficiency and to reduce rhizosphere microbial growth and pathogen attack, (3) direct uptake to recapture organic nutrients released from soil organic matter, and (4) for inter-root and root–microbial signal exchange. Due to severe flaws in the interpretation of commonly used isotopic labelling techniques, there is still great uncertainty surrounding the importance of these individual fluxes in the rhizosphere. Due to the importance of rhizodeposition in regulating ecosystem functioning, it is critical that future research focuses on resolving the quantitative importance of the different C and N fluxes operating in the rhizosphere and the ways in which these vary spatially and temporally.     

不同林龄杉木人工林菌根侵染特征研究

丛枝菌根真菌是一种通过土壤侵染植物根系,与寄主植物互利共生的重要有益真菌。探究不同林龄杉木林中菌根侵染状况与土壤养分的变化规律,有利于深入认识丛枝菌根真菌—杉木相互作用的养分调控因素,从而为改善杉木人工林土壤肥力、促进杉木林可持续经营提供依据。分别选取10a、25a、45a杉木纯林,分析了不同林分菌根侵染率与孢子密度及部分土壤养分因子(全磷、速效磷、全钾、速效钾)的关系。结果表明:(1)菌根侵染率与孢子密度均呈现出随林龄增大而增大的趋势,pH随林龄增大而减少的趋势;(2)根际土中磷的含量总体偏低,而且受到土壤酸化流失和丛枝菌根真菌积累的双重影响呈现出先减少后增加的趋势;(3)虽然有效钾含量随林龄变化趋势不显著,但丛枝菌根真菌能促进土壤钾的积累。因此,丛枝菌根真菌能有效调控根际土的养分动力学特征,减缓土壤酸化造成的养分流失。     

Root uptake and translocation of radiocaesium from agricultural soils by various plant species

Plant uptake of radiocaesium from soil is an important pathway for the entry of this pollutant into the human food chain and so contributes to any assessment of the radiation dose following contamination. Large differences in soil–plant transfer factors have been reported for plant species grown on the same soils. Few studies have attempted to distinguish between differences in root uptake and root-to-shoot translocation. We have investigated the root uptake of radiocaesium from artificially contaminated soils and the subsequent translocation to shoots for various plant species grown on three agricultural soils. The effects of short contact times and potassium starvation or enrichment have been studied. The Cs adsorption properties of rhizosphere soils have been compared with those of the initial soils. The proportion of activity removed from soil is largely soil dependent. Root uptake properties have less effect, but appear to be species determined, and not influenced by soil properties. Differences in soil-to-shoot transfer factor arise from species-dependent differences in root-to-shoot translocation. Root-to-shoot activity ratios are not soil dependent. There was little effect of soil potassium status. Root action slightly enhanced Cs adsorption on one soil, probably due to mineral weathering associated with the release of nonexchangeable potassium.     

Plant species response to variation in soil nutrient saturation gradients in Zaraninge Forest,Tanzania

Zaraninge is a former forest reserve which is now included as part of the Saadani National Park in Bagamoyo District Coast Region. The influence of soil mineral nutrients (N, P, K, Mg and Na) on the pattern of plant species distributions was determined, and transect method was used for data collection. Plant species composition and abundance followed variation in soil nutrient gradients within Zaraninge Forest. Canonical correspondence analysis of soil and plant data showed that calcium, magnesium and phosphorus were the mineral nutrients that significantly influenced the distribution plant species (P < 0.05) while other nutrients had no significant effects. Data from this study provide evidence of the existing variation in soil nutrient saturation gradients to which plant species distribution responds within the forest. A successful conservation of Zaraninge Forest biodiversity requires protecting the soil which is the necessary habitat for the plant species.     

Effects of ammonium sulphate application on the rhizosphere,fine-root and needle chemistry in aPicea abies (L.) Karst. stand

Rhizosphere, fine-root and needle chemistry were investigated in a 28 year old Norway spruce stand in SW Sweden. The uptake and allocation pattern of plant nutrients and aluminium in control plots (C) and plots repeatedly treated with ammonium sulphate (NS) were compared. Treatments started in 1988. Current year needles, one-year-old needles and cylindrical core samples of the LFH-layer and the mineral soil layers were sampled in 1988, 1989 and 1990. Compared to the control plots, pH decreased significantly in the rhizosphere soil in the NS plots in 1989 and 1990 while the SO₄-S concentration increased significantly. Aluminium concentration in the rhizosphere soil was generally higher in the NS plots in all soil layers, except at 0–10 cm depths, both in 1989 and 1990. Calcium, Mg and K concentrations also increased after treatment with ammonium sulphate. Ammonium ions may have replaced these elements in the soil organic matter. The NS treatment significantly reduced Mg concentrations in fine roots in all layers in 1990. A similar trend was found in the needles. Ca concentrations in fine roots were significantly lower in the NS plots in the LFH layer in 1990 and the same pattern was found in the current needles. The N and S concentrations of both fine roots and needles were significantly higher in the NS plots. It was suggested that NS treatment resulted in displacement of Mg, Ca and K from exchange sites in the LFH layer leading to leaching of these cations to the mineral soil. Further application of ammonium sulphate may damage the fine roots and consequently adversely affect the water and nutrient uptake of root systems.     

Effects of hydrology on spatial patterns of soil development in created riparian wetlands

Surficial soil development was studied in four wetland basins created on the floodplain of the Des Plaines River near Chicago, Illinois, USA. These studies determined changes in the spatial distribution of plant-available nutrients as a result of establishing two different wetland hydrologic regimes. Three wetland basins had mineral soils and one an organic soil. A geostatistical analysis including kriging of collected data indicated that all soil parameters showed significant changes in their spatial structure as a result of the water inputs and unidirectional flows. The degree of spatial variability as indicated by autocorrelation in the soil data (i.e., points closer to one another are more similar than points further apart due to the influence of landscape processes) declined for all parameters except Mg⁺². Temporal changes in the spatial patterns of extractable phosphorus (P) and percent organic carbon (OC) tended to be inverse; P declined in areas where OC increased and vice versa. The spatial pattern of these changes was dissimilar in the mineral soils as compared to the organic soil and was related to patterns of primary productivity. Zones of P uptake and OC accumulation were also related to wetland hydrology and primary productivity. Changes in the distribution of nutrients, particularly P, may be viewed as a result of nutrient spirals within the wetlands. By comparison, the reorganization in the concentrations of K⁺ and Ca⁺² appear to have been mediated by cation exchange processes. The formation of new concentration gradients was strongly related to both flow pathways and the different water inflow rates. The formation of concentration gradients in exchangeable cations was not reflected in the average concentrations within each basin. Mean values changed significantly in only a few instances. Reducing data in this way missed important biogeochemical changes occurring within the experimental wetland basins. This revised version was published online in July 2006 with corrections to the Cover Date.     

Relative contributions of soil chemistry, plant physiology and rhizosphere induced changes in speciation on Ni accumulation in plant shoots

The aim of this study is to rank the relative importance of soil properties, root uptake and root-to-shoot redistribution on the transfer of the trace element nickel from soil to the shoots of non hyperaccumulatings plants. Two contrasting soils and seven plant species have been studied using the radioactive isotope, ⁶³Ni. Shoots and roots were analysed separately and the specific activity of each plant has been measured. The isotopic exchange properties of rhizosphere soil where compared with control non rhizosphere soil. Possible changes in Ni speciation in the rhizosphere have been assessed by comparing the isotopic exchange properties of the rhizosphere and control soil and by comparing the specific activities of Ni in each plant. The capacity of soil to immobilise added radiotracer largely determines root uptake, leading to between a 4- and 40-fold difference between soils for a given species. The redistribution of nickel from roots to shoots was fairly constant for plants grown on the rendzina, but varied strongly between species for the acid soil. This variation enhanced the contrast between species of the soil-to-shoot transfer factor. Root action significantly enhanced immobilisation of added nickel in an acid soil due to the modification of speciation of initially non exchangeable soil nickel, but had little effect on a neutral rendzina. Changes in rhizosphere pH were similar on the two soils. In the acid soil, these pH changes were accompanied by changes in Ni speciation but a causative link has not been established. In the neutral soil pH changes may have modified root uptake properties.     

Root Development and Nutrient Uptake

Root system formation proceeds in close coordination with shoot growth. Accordingly, root growth and its functions are regulated tightly by the shoot through materials cycling between roots and shoots. A plant root system consists of different kinds of roots that differ in morphology and functions. The spatial configuration and distribution of these roots determine root system architecture in the soil, which in turn primarily regulates the acquisition of soil resources like nutrients and water. Morphological and physiological properties of each root and the concomitant tissues further affect nutrient uptake and transport, while the root traits that are related to such acquisition also depend on the kinds of nutrients and their mobility in the soil. In addition, mechanisms involved in the uptake and transport of mineral nutrients recently have been elucidated at the molecular level. A number of genes for acquisition and transport of various mineral nutrients have been identified in model plant systems such as Arabidopsis thaliana, and rice, and in other plant species. An integration of studies on nutrient behavior in soils and the morphological and physiological functions of root systems will further elucidate the mechanism of plant nutrient uptake and transport by roots, and offer a real possibility of genetically improving crop productivity in problem soils.