A mathematical model of phytoremediation for petroleum contaminated soil: sensitivity analysis

Phytoremediation is an attractive treatment technology for many contaminated sites due to its cost effectiveness and public acceptance. We present a sensitivity analysis of important parameters from a screening level model for phytoremediation by grass species of weathered petroleum-contaminated sites. The conceptual framework is that root movement through contaminated soil will enhance contaminant biodegradation by providing a local environment more favorable for petroleum degrading microorganisms—the so-called rhizosphere effect. Common questions in phytoremediation are, “What species should be planted?” and “What management practices should be followed?” These choices may affect degradation kinetics, root biomass (and therefore rhizosphere volume), and the root turnover. Important model parameters are the rate constants, rhizosphere volume, and the rate of root turnover. We present a sensitivity analysis with the aim of identifying the most important factors for improving phytoremediation effectiveness. For simulations of the phytoremediation of weathered diesel range organics, our results indicate that annual species, with higher root turnover, are preferred over perennial species with the caveat of equal degradation rate constants, that is, no species-dependent effects. In addition, the results suggest that the management of nonrhizosphere soil could play an important role in the overall effectiveness of phytoremediation. Finally, the effect of increasing root biomass or increasing the rhizosphere thickness is approximately equivalent with respect to the ultimate removal of the contaminants.     

郭岩山不同海拔丝栗栲细根功能性状及其与土壤因子的关系

为揭示丝栗栲(Castanopsis fargesii)细根功能性状对环境变化的适应机制,对郭岩山500、700、900 m海拔处丝栗栲细根功能性状及其与土壤因子的关系进行研究。结果表明,丝栗栲细根生物量与细根根长密度、表面积密度、组织密度及体积密度呈正相关,细根根长密度、体积密度、表面积密度和比根长4个性状间均呈极显著正相关关系,且均与细根组织密度呈显著负相关。根际土含水量、C和N含量与细根比根长、根长密度、体积密度、表面积密度均存在显著正相关关系,而土壤容重与细根组织密度呈正相关。海拔700 m的细根生物量、根长密度、表面积密度及体积密度显著大于海拔500和900 m的。500和900 m海拔的根长密度、表面积密度与土壤深度呈负相关,而500 m海拔细根的组织密度与土壤深度呈正相关。因此,郭岩山丝栗栲通过改变细根功能性状来适应海拔和土壤的变化。     

杉木成熟林细根形态与功能特征的海拔梯度变异特点

为探究植物对环境变化的适应策略,在安徽省金寨县天马国家自然保护区,以不同海拔高度(750、850、1000、1150 m)杉木(Cunninghamia lanceolata)成熟林为对象,采用土钻法获取土壤细根样品,分别测定了不同海拔不同土层(0—10 cm、10—20 cm、20—30 cm)土壤细根生物量、形态特征参数和碳氮含量。结果表明:(1)随海拔梯度增加,0—30 cm土层细根生物量、根长密度、比根长、表面积密度、体积密度均呈先减少后增加趋势,在海拔750 m生物量最大,其余指标在海拔1150 m最大;随土层深度增加,同一海拔细根生物量、根长密度、表面积密度、体积密度均呈减少趋势。(2)随海拔梯度增加,0—30 cm土层细根C和N含量呈先增加后减少趋势,C/N比呈先减少后增加再减少趋势;随土层深度增加,同一海拔细根C含量呈先减少后增加趋势,N含量呈降低趋势,C/N比呈上升趋势。(3)细根N含量与生物量、根长密度和体积密度显著正相关,C/N比与生物量、根长密度、表面积密度和体积密度极显著负相关。(4)土壤水分对细根生物量及其形态指标影响显著。     

根区交替控制灌溉条件下玉米根系吸水规律

为了阐明根区交替控制灌溉(CRDAI)条件下玉米根系吸水规律,通过田间试验,在沟灌垄植模式下采用根区交替控制灌溉研究玉米根区不同点位(沟位、坡位和垄位)的根长密度(RLD)及根系吸水动态。研究表明,根区土壤水分的干湿交替引起玉米RLD的空间动态变化,在垄位两侧不对称分布,并存在层间差异;土壤水分和RLD是根区交替控制灌溉下根系吸水速率的主要限制因素。在同一土层,根系吸水贡献率以垄位最大,沟位最低;玉米营养生长阶段,10—30 cm土层的根系吸水速率最大;玉米生殖生长阶段,20—70 cm为根系吸水速率最大的土层,根系吸水贡献率为43.21%—55.48%。研究阐明了交替控制灌溉下根系吸水与土壤水分、RLD间相互作用的动态规律,对控制灌溉下水分调控机理研究具有理论意义。     

青杨人工林根系生物量、表面积和根长密度变化

在植物生长季节,采用钻取土芯法对秦岭北坡50年生青杨人工林根径≤2 mm和2～5 mm根系的生物量、表面积和根长密度进行测定.结果表明：在青杨人工林根系(＜5 mm)中,根径≤2 mm根系占总生物量的77.8%,2～5 mm根系仅占22.2%;根径≤2 mm根系表面积和根长密度占根系总量的97%以上,而根径2～5 mm根系不足3%.随着土层的加深,根径≤2 mm根系生物量、表面积和根长密度数量减少,根径2～5 mm根系生物量、表面积和根长密度最小值均分布在20～30 cm土层.≤2 mm根系生物量、表面积和根长密度与土壤有机质、有效氮呈极显著相关,而根径2～5 mm根系的相关性不显著.     

漓江水陆交错带典型立地根系分布与土壤性质的关系

研究根系与土壤关系是发掘河岸带生态退化等问题内在原因的重要途径。在漓江流域水陆交错带选取缓坡、陡坡、江心洲、人工岸坡4种典型立地类型,对不同土层深度的根长密度、根系生物量、比根长,以及根系特征与土壤有机质、全氮、有效磷的关系进行了研究,旨在为漓江流域生态修复过程中植被恢复、植被配置、快速绿化材料选取提供科学依据。结果表明:(1)同一立地类型0—10 cm土层和10—20 cm土层比根长差异性不显著。0—10 cm到10—20 cm土层,各立地类型根长密度和根系生物量密度均减小,但不同立地类型根长密度和根系生物量密度的差异程度逐渐缩小,表明地形、地表植物类型及生长状况对根长密度分布的影响也随土层深度的增加而逐渐减小。细根根长和生物量随着土壤深度的增加而减小。(2)土壤有机质含量差异性显著,分布规律为人工岸坡陡坡江心洲缓坡;土壤全氮含量从大到小依次是人工岸坡、陡坡、缓坡、江心洲,其值分别为:3.12、2.33、1.56、1.32 g/kg;土壤全氮与土壤有机质呈显著正相关。江心洲和缓坡有效磷含量远远大于人工岸坡和陡坡,原因是漓江水长期受人为洗漱影响,导致受江水干扰大的立地类型有效磷含量高。(3)根长密度、比根长、根系生物量与有机质、全氮含量呈正相关,与有效磷含量呈负相关,说明土壤根系越丰富,越有利于增加土壤有机质和全氮含量,但遏制了土壤有效磷。细根长度、生物量与根长密度在0.01水平(双侧)上显著正相关,与根系生物量密度呈负相关。     

Modelling the effect of varying soil water on root growth dynamics of annual crops

We present a simple framework for modelling root growth and distribution with depth under varying soil water conditions. The framework considers the lateral growth of roots (proliferation) and the vertical extension of roots (root front velocity). The root front velocity is assumed to be constant when the roots descend into an initially wet soil profile. The lateral growth of roots is governed by two factors: (1) the current root mass or root length density at a given depth, and (2) soil water availability at that depth.Under non-limiting soil water conditions, the increase in root mass at any depth is governed by a logistic equation so that the root length density (R _v) cannot exceed the maximum value. The maximumR _v, is assumed to be the same for all depths. Additional dry matter partitioned to roots is initially distributed according to the current root mass at each depth. As the root mass approaches the maximum value, less dry matter is partitioned to that depth.When soil water is limiting, a water deficit factor is introduced to further modify the distribution of root dry matter. It is assumed that the plant is an energy minimiser so that more root mass is partitioned to the wetter regions of the soil where least energy will be expended for root growth. Hence, the model allows for enhanced root growth in areas where soil water is more easily available.Simulation results show that a variety of root distribution patterns can be reproduced due to varying soil water conditions. It has been demonstrated that broad patterns of root distribution reported in the literature can also be simulated by the model.     

Fine‐root exploitation strategies differ in tropical old growth and logged‐over forests in Ghana

Understanding the changes in root exploitation strategies during post‐logging recovery is important for predicting forest productivity and carbon dynamics in tropical forests. We sampled fine (diameter < 2 mm) roots using the soil core method to quantify fine‐root biomass and architectural and morphological traits to determine root exploitation strategies in an old growth forest and in a 54‐yr‐old logged‐over forest influenced by similar parent material and climate. Seven root traits were considered: four associated with resource exploitation potential or an ‘extensive’ strategy (fine‐root biomass, length, surface area, and volume), and three traits which reflect exploitation efficiency or an ‘intensive’ strategy (specific root area, specific root length, and root tissue density). We found that total fine‐root biomass, length, surface area, volume, and fine‐root tissue density were higher in the logged‐over forest, whereas the old growth forest had higher total specific root length and specific root surface area than the logged‐over forest. The results suggest different root exploitation strategies between the forests. Plants in the old growth forest invest root biomass more efficiently to maximize soil volume explored, whereas plants in the logged‐over forest increase the spatial distribution of roots resulting in the expansion of the rhizosphere.     

弓杠岭不同海拔云杉细根生物量及形态特征

海拔变化是多环境因子的梯度效应,细根作为植物吸收水分与养分的重要器官,其性状特征在指示植物的生长和分布等方面意义重大.该研究以弓杠岭2500～3300 m海拔地的云杉(Picea asperata)细根为研究对象,采用根序分级法对云杉1～5级根序的生物量及细根形态(平均直径、比根长、根长密度、比表面积)进行测定,以明确...     

Field performance of Solanum sisymbriifolium, a trap crop for potato cyst nematodes. II. Root characteristics

Hatching of potato cyst nematodes is induced by root exudates of Solanaceae, such as Solanum sisymbriifolium, and is therefore related to root length distribution of this crop. A mathematical model was derived to relate the hatching potential to root length density (RLD). A series of field experiments was carried out to study actual root length distribution of S. sisymbriifolium in relation to shoot properties and to provide input into the model. Using a modified Poisson distribution formula for the three‐dimensional distribution of roots in a volume of soil, the relation between the zone of influence of hatching agents and the RLD could be derived. On this basis, the minimal RLD was estimated, which is needed to expose 75%, 90% or 95% of cysts to root exudates, as a function of the length of the zone of influence of hatching agents on cysts. The logarithm of the total root length showed a linear relation with the logarithms of above‐ground biomass and with leaf area index. Root diameter distribution was the same for all crops examined and independent of soil depth. Fine roots (<0.4 mm in diameter) constituted around 50% of total root length. Using a zone of influence of 1.00, 0.75 and 0.50 cm around the centre of each root, a minimal RLD for sufficient soil exploration (75%) was estimated. Depth at which that minimal RLD was exceeded was linearly related to total root length (km m^?2) and to above‐ground crop biomass, enabling estimations being made of the potential hatching efficacy as related to measurable properties of S. sisymbriifolium crops. The proposed approach to derive potential hatching effects from crop properties needs further validation; particularly, the distance of influence of root exudates is a critical factor.     

毛竹种群向针阔林扩张的根系形态可塑性

为了弄清毛竹(Phyllostachys edulis)向针阔林扩张过程中根系的形态可塑性反应,在浙江天目山自然保护区毛竹向针阔林扩张的典型过渡地带,连续区域上设置毛竹纯林、针阔-毛竹混交林(以下简称过渡林)、针阔林3种样地。用根钻法采集样地毛竹根系、针阔树根系并比对其生物量密度、细根比根长、相邻同级侧根节点距等形态特征参数变化。结果表明:随着毛竹的扩张程度增加,林内根系生物量密度增加;且与针阔树竞争过程中毛竹将更多的根系放置于表层;同时在水平方向上随离样株距离的增加未出现明显变化,而针阔树根系则随离样木距离的增加而逐渐减少;毛竹根系比根长明显增加,平均增幅15%;一、二级侧根节点距则均有所下降,毛竹侧根数量增多。这些结果表明毛竹种群可通过根系生物量密度、细根比根长、相邻同级侧根节点距等形态可塑性方式实现向周边森林扩张。     

Theoretical evidence that root penetration ability interacts with soil compaction regimes to affect nitrate capture

Background and AimsAlthough root penetration of strong soils has been intensively studied at the scale of individual root axes, interactions between soil physical properties and soil foraging by whole plants are less clear. Here we investigate how variation in the penetration ability of distinct root classes and bulk density profiles common to real-world soils interact to affect soil foraging strategies.MethodsWe utilize the functional–structural plant model ‘OpenSimRoot’ to simulate the growth of maize (Zea mays) root systems with variable penetration ability of axial and lateral roots in soils with (1) uniform bulk density, (2) plow pans and (3) increasing bulk density with depth. We also modify the availability and leaching of nitrate to uncover reciprocal interactions between these factors and the capture of mobile resources.Key ResultsSoils with plow pans and bulk density gradients affected overall size, distribution and carbon costs of the root system. Soils with high bulk density at depth impeded rooting depth and reduced leaching of nitrate, thereby improving the coincidence of nitrogen and root length. While increasing penetration ability of either axial or lateral root classes produced root systems of comparable net length, improved penetration of axial roots increased allocation of root length in deeper soil, thereby amplifying N acquisition and shoot biomass. Although enhanced penetration ability of both root classes was associated with greater root system carbon costs, the benefit to plant fitness from improved soil exploration and resource capture offset these.ConclusionsWhile lateral roots comprise the bulk of root length, axial roots function as a scaffold determining the distribution of these laterals. In soils with high soil strength and leaching, root systems with enhanced penetration ability of axial roots have greater distribution of root length at depth, thereby improving capture of mobile resources.     

Competition in tree row agroforestry systems. 1. Distribution and dynamics of fine root length and biomass

Complementarity in the distribution of tree and crop root systems is important to minimise competition for resources whilst maximising resource use in agroforestry systems. A field study was conducted on a kaolinitic Oxisol in the sub-humid highlands of western Kenya to compare the distribution and dynamics of root length and biomass of a 3-year-old Grevillea robusta A. Cunn. ex R. Br. (grevillea) tree row and a 3-year-old Senna spectabilis DC. (senna) hedgerow grown with Zea mays L. (maize). Tree roots were sampled to a 300 cm depth and 525 cm distance from the tree rows, both before and after maize cropping. Maize roots were sampled at two distances from the tree rows (75–150 cm and 450–525 cm) to a maximum depth of 180 cm, at three developmental stages. The mean root length density (L_rv) of the trees in the upper 15 cm was 0.55 cm cm⁻³ for grevillea and 1.44 cm cm⁻³ for senna, at the start of the cropping season. The L_rv of senna decreased at every depth during the cropping season, whereas the L_rv of grevillea only decreased in the crop rooting zone. The fine root length of the trees decreased by about 35% for grevillea and 65% for senna, because of maize competition, manual weeding, seasonal senescence or pruning regime (senna). At anthesis, the L_rv of maize in the upper 15 cm was between 0.8 and 1.5 cm cm⁻³. Maize root length decreased with greater proximity to the tree rows, potentially reducing its ability to compete for soil resources. However, the specific root length (m g⁻¹) of maize was about twice that of the trees, so may have had a competitive uptake advantage even when tree root length was greater. Differences in maize fine root length and biomass suggest that competition for soil resources and hence fine root length may have been more important for maize grown with senna than grevillea. This revised version was published online in June 2006 with corrections to the Cover Date.     

瓦屋山常绿阔叶次生林建群种扁刺栲的细根分布及其碳氮特征

四川瓦屋山国家森林公园是我国西部的中亚热带湿性常绿阔叶林的典型代表,具有四川现存的较为完好的扁刺栲(Castanopsis platyacantha)-华木荷(Schima sinensis)群系,该研究利用土钻法探讨了该群系内主要建群种扁刺栲标准木的细根分布及其碳氮特征。结果表明:(1)扁刺栲细根总生物量为173.62 g·m~(-2),其中活根生物量为135.29 g·m~(-2)。(2)随着土层深度的增加,扁刺栲细根生物量、根长密度、根系表面积和比根长呈下降趋势,0~30 cm土层所占比例分别为67.23%、69.53%、69.48%和57.20%;根长密度、根系表面积和比根长均随细根直径的增加而显著下降,直径小于1 mm的根系所占比例分别为58.84%、52.59%和51.36%。(3)扁刺栲细根生物量、根长和表面积消弱系数β均随根系直径的增加而增加。(4)根系C含量在第Ⅰ土层中随细根直径的增加而显著增加,在其他土层则无显著差异;直径小于2 mm的根系C含量在第Ⅰ土层中显著低于其他土层,大于2 mm的根系C含量在各土层间的差异不大。(5)根系N含量随根系直径和土层深度的增加而减少,C/N值则与之相反。该研究结果在一定程度上反映了该次生林地下细根的垂直分布及养分特征,为揭示该生态系统地下生态过程及今后在该生态系统研究环境变化对地下生态过程的影响提供了基础数据。     

翅荚木人工林不同径阶间细根主要功能性状与根际土壤养分的关系

细根能敏感地感知土壤环境变化,对植物生长发育具有重要影响.以6年生翅荚木人工林为对象,对其不同径阶的细根主要功能性状与根际土壤养分特征及两者间关系进行分析.结果表明:细根生物量、根长密度与根体积密度均随径阶增加而增加,比根长与比根面积则随径阶增加呈先升高再下降后升高的趋势,根组织密度则与径阶大小不相关.不同径阶翅荚木根际土壤的pH值及含水率、全碳、全磷、铵态氮、硝态氮和总有效氮含量均存在显著差异,大径阶林木的根际土壤全碳、全氮、硝态氮、总有效氮含量相对较高,小径阶林木的根际土壤含水率、土壤全磷、铵态氮含量相对较高.土壤全氮、全碳、硝态氮和总有效氮含量与林木细根的生物量、根长密度、根体积密度呈显著正相关;土壤全磷与林木细根的根组织密度呈显著正相关,与比根长、比根面积呈显著负相关;土壤含水率与林木细根的生物量和根体积密度均呈显著正相关;根际土壤pH和林木细根的比根长、比根面积呈显著正相关,与根组织密度则呈显著负相关.研究结果可为翅荚木优良种质资源选育提供科学依据.     

基于森林调查数据的长白山天然林森林生物量相容性模型

森林生物量估算是进行陆地生态系统碳循环和碳动态分析的基础,但现有估测模型存在着总量与分量不相容的问题.本文以吉林省汪清天然林区为例,提出了基于森林调查的相容性森林生物量模型设计思想,并采用联立方程组为不同森林群落构造了一系列引入林分蓄积因子的相容性生物量模型,得到的预估精度较高.其中,针叶林、阔叶林和针阔混交林群落的森林生物量模型预估精度均在95%以上,基本上解决了森林生物量模型的相容性问题.     

Effects of water supply on root traits and biological yield of Durum (Triticum durum Desf.) and Khorasan (Triticum turanicum Jakubz) wheat

The ability of a plant to change its root characteristics to increase the acquisition of soil water is an important adaptation mechanism to water limitation. In this regard, a field study was carried out in the Pannonian region of eastern Austria with two tetraploid wheat genotypes, i.e. Durum (Triticum durum Desf.) and Khorasan (Triticum turanicum Jakubz), during a comparatively wetter and drier year, i.e. 2008 and 2009, respectively. The genotypes showed significant differences in average root diameter and fine root length. All root traits varied with soil depth. The highest root length density and root biomass were observed with Khorasan wheat in 0–10 cm soil depth. Durum wheat showed a stronger response in fine roots to water availability and produced more fine roots in the moist year. Electric root capacitance was higher with Khorasan wheat. Durum showed higher biological yield stability across years with different precipitation with respect to above- and belowground biomass. It produced more leaf area under humid conditions. Khorasan allocated more assimilates to belowground organs in dry conditions, but without positive effect on aboveground biomass.     

黄土高原白羊草、沙棘和辽东栎细根比根长特性

以黄土高原地区典型草本（白羊草）、灌木（沙棘）和乔木（辽东栎）为对象,研究了3种植物细根比根长在不同土层的分布状况以及与其它细根参数和土壤物理因子之间的相关性。结果表明,3种植物细根比根长的变化范围为6～55ram／rag。在0,80cm土层,白羊草、沙棘和辽东栎细根比根长变化范围分别为18—55mm／mg,14—4JDmm／mg,6—33mm／mg。3种植物0--80cm土层平均细根比根长从大到小依次为白羊草〉沙棘〉辽东栎。3种植物0-10cm土层细根比根长依次为沙棘〉辽东栎〉白羊草,10-80cm依次为白羊草〉辽东栎〉沙棘,表明3种植物细根比根长不仅在这两土层中的分布不具一致性,而且与0-80cm土层平均比根长也不具有一致性,进一步说明3种植物沿土壤剖面的生物量分配策略不同。相关分析表明,3种植物细根比根长与其它细根参数之间的相互关系各不相同,制约程度存在差异。与土壤物理因子的相关分析表明,3种植物细根比根长均随土壤含水量的增加而减少。土壤各级水稳性团聚体和土壤颗粒对3种植物细根比根长并无一致的影响。     

Fine roots, arbuscular mycorrhizal hyphae and soil nutrients in four neotropical rain forests: patterns across large geographic distances

* It is commonly hypothesized that stand-level fine root biomass increases as soil fertility decreases both within and among tropical forests, but few data exist to test this prediction across broad geographic scales. This study investigated the relationships among fine roots, arbuscular mycorrhizal (AM) fungi and soil nutrients in four lowland, neotropical rainforests. * Within each forest, samples were collected from plots that differed in fertility and above-ground biomass, and fine roots, AM hyphae and total soil nutrients were measured. * Among sites, total fine root mass varied by a factor of three, from 237+/-19 g m-2 in Costa Rica to 800+/-116 g m-2 in Brazil (0-40 cm depth). Both root mass and length were negatively correlated to soil nitrogen and phosphorus, but AM hyphae were not related to nutrients, root properties or above-ground biomass. * These results suggest that understanding how soil fertility affects fine roots is an additional factor that may improve the representation of root functions in global biogeochemical models or biome-wide averages of root properties in tropical forests.     

Effects of liming on rhizosphere chemistry and growth of fine roots and of shoots of sessile oak (Quercus petraea)

Soil acidification can be detrimental to root growth and nutrient uptake, and liming may alleviate such acidification. In the following study, seedlings of sessile oak (Quercus petraea Liebl. M.) were grown in rhizotrons and subjected to liming (L) or gypsum (G) treatments and compared with the control (C). In order to study and interpret the impact of these calcium rich treatments on fine root development and tree growth, the following parameters were assessed: fine root biomass, fine root length, seedling development (height, diameter, leaves), seedling biomass, nutrient content of roots and seedlings, bulk soil and soil solution chemistry and rhizosphere soil chemistry. The results show that liming increased bulk soil pH, exchangeable Mg, Ca and the Ca/Al molar ratio, and decreased exchangeable Al, mainly in the A-horizon. Gypsum had a similar but smaller impact on exchangeable Al, Ca, H⁺ and the Ca/Al molar ratio in the A-horizon, but reacted with depth, so that exchangeable Mn, Mg and Ca were increased in the B-horizon. In the rhizosphere, the general pattern was determined by the treatment effects of the bulk soil. Most elements were more concentrated in the rhizosphere than in bulk soil, except for Ca which was less concentrated after liming or gypsum application. In the B-horizon rhizosphere pH was increased by the treatments (L > G,C) close to the root tips. Furthermore, the length of the zone with a positive root-induced pH increase was greater for the limed roots as compared with both the other treatments. Fine root growth was stimulated by liming (L > G,C) both in terms of biomass and length, whereas specific root length was not obviously affected apart from the indication of some stimulation after liming at the beginning. The live:dead ratio of fine roots was significantly higher in the limed rhizotrons as compared to the control (G not assessed), indicating lower mortality (higher longevity). Shoot growth showed greater lime-induced stimulation (L > G,C) as compared to root growth. As a result the shoot:root ratio was higher in the limed rhizotrons than in the control (L > G,C). Liming induced a higher allocation of P, S, Mg, Ca and K to the leaves, stem and twigs. Gypsum showed similar effects, but was only significant for S. Liming increased the foliar Ca/Al ratio by both increasing foliar Ca and decreasing foliar Al, whereas gypsum did not clearly improve foliar nutrition. This study suggests that a moderate application of lime can be successful in stimulating seedling growth, but that gypsum had no effect on seedling growth. It can be concluded that this lime-induced growth stimulation is directly related to the improved soil fertility status, and the alleviation of Al toxicity and acid stress, resulting in better foliar nutrition. The impact of liming on fine roots, as a consequence, was not limited to a stimulation of the total amount of fine roots, but also improved the root uptake performance. This revised version was published online in June 2006 with corrections to the Cover Date.