Wetting and drying cycles in the maize rhizosphere under controlled conditions. Mechanics of the root-adhering soil

Mechanical properties of the topsoil (sandy Podsol and silty Luvisol, FAO) adhering to maize (Zea mays L.) roots and its bulk soil counterpart were studied as a function of soil texture and final soil water suction at harvest, with three soil water suction values of approximately 30, 50 and 60 kPa. Two scales of observation were also selected: the whole soil:root system and the root-adhering soil aggregates. Three methods were used to characterize the stability of the soil:root system: mechanical shaking in air, and dispersion by low-power ultrasonication, with or without preliminary immersion of the soil:root system in water. Soil disruption kinetics, which were fitted with first-order kinetics equations, were analyzed and discussed. For example, silty soil ultrasonication kinetics, without preliminary water-immersion, could be divided into two parts: the first faster part, which was characterized by a mean rate K value of 6.8–7.2 mJ^-1, is attributed to soil slaking, whereas the second slower part, which was characterized by a mean rate K value of 1.5–1.6 mJ^-1, was attributed to the rupture of the `firmly root-adhering soil' from the roots. A clear plant effect was observed for both aggregate tensile strength and friability, with higher aggregate strength for the root-adhering silty soil (450–500 kPa) than for its bulk silty soil counterpart (410–420 kPa), and lower friability (coefficient of variation of the aggregate strength) for the root-adhering silty soil (e.g. 67% at a soil water suction value of 30 kPa) than for its bulk silty soil counterpart (e.g. 49% at asoil water suction value of 30 kPa). These effects were attributed to root exudation, which was significantly higher for the driest silty topsoil than for the wetter ones. In conclusion, the mechanical properties of the silty topsoil adhering to the maize roots are attributed to both physical and biological interactions occurring in the maize rhizosphere. This revised version was published online in June 2006 with corrections to the Cover Date.     

The deformation of soil by penetrometers and root tips ofPisum sativum

Channels were formed by seminal roots ofPisum sativum and a steel penetrometer of similar dimeter in blocks of remoulded and weathered soil. For both types of channels, the soil was equilibrated and maintained at –12kPa matric water potential during formation. Small samples of soil containing channels were then excavated and examined using a scanning electron microscope. Sections of root channels were found to contain a clearly differentiated zone of newly remoulded soil containing oriented clay. In contrast to channels created by the rigid steel probe, the newly remoulded zone surrounding root channels did not exhibit either a region of maximum soil compression at the channel surface or a radial pattern of shear failure and compression. This micromorphological evidence suggests that exudates may have an additional role to play in reducing the mechanical strength of soil in the proximity of the root tip. The mechanism is thought to operate through an accumulation of soil water related to solute potential and a resultant increase in matric potential.     

A comparison of penetrometer pressures and the pressures exerted by roots

Summary Previous work is reviewed in which the ratio of the pressures required for soil penetration by roots and penetrometers are compared. It appears that this ratio can vary from about 2 to 8 depending on conditions. However, there is very little experimental evidence and most of the work has been inferential.Direct measurements are reported for the stresses exerted by a 1 mm diameter penetrometer probe and by the roots of pea seedlings when penetrating Urrbrae fine sandy loam. Six soil conditions were used: (non-weathered remoulded soil cores + artificially weathered remoulded soil cores + undisturbed field clods) × (confined + unconfined cores or clods). The confinement treatment was to test for any effects of additional restraint to cylindrical root expansion. The weathering and field clod treatments were to test the hypothesis that root elongation is facilitated by tensile failure ahead of the root tip.The principal conclusions are as follows. The laboratory weathering treatment reduced the soil tensile strength by 25%. This resulted in a small but significant reduction in the pressure for root penetration into confined cores. Compared with remoulded non-weathered cores, field clods had a 2 to 3 fold greater penetrometer resistance and a 50% lower tensile strength. The force required for root penetration into unconfined field clods was only 10% greater than for unconfined non-weathered cores. For the former (which is closest to field conditions) the penetrometer had to exert a pressure 5.1 times greater than a root tip in order to penetrate the soil. Penetrometer penetration pressure was independent of probe diameter in the 1–2 mm range in the soil used. Core confinement restricts root radial expansion and modifies the penetration force of metal probes and plant roots.On the basis of the new results it is tentatively concluded that soil tensile failure can facilitate penetration by roots.     

植物根系固坡抗蚀的效应与机理研究进展

植物根系对抵抗坡体浅层滑坡和表土侵蚀起着巨大的作用.植物根系通过增强土体的抗剪强度发挥固坡效应.目前有关植物根系固坡机理的模型较多,普遍接受的是Wu-Waldron模型.该模型表明,植物根系产生的土体抗剪强度的增量与根系的平均抗拉强度和根面积比成正比,应用该模型评价根系固坡效应的2个最重要因素是根系的平均抗拉强度和根面积比.研究发现,土壤抗侵蚀性随着植物根系数量的增加而提高,但未有一致的定量函数关系.植物根系提高土壤抗侵蚀性主要通过直径小于1mm的须根起作用.须根通过增加土壤水稳性团聚体的数量与粒径等作用来提高土壤的稳定性,以抵抗水流分散;须根还能有效地增强土壤渗透性,减少径流,从而达到减少土壤冲刷的目的.     

Soil Anti-Scouribility Enhanced by Plant Roots

The magnitude of soil anti-scouribility depends on the physical condition of the soil. Plant roots can greatly enhance soil stability and anti-erodibility. A scouring experiment of undisturbed soil was conducted to investigate the effects of roots on soil anti-scouribility and its distribution in the soil profile. At the end of each erosion test, plant roots were collected from soil samples and root surface area was calculated by means of a computer image analysis system (CIAS). Root surface area density (RSAD), the surface area of the roots per unit of soil volume, was related to soil anti-scouribility. More than 83% of root surface area was concentrated in the 0-30 cm soil layer. Soil anti-scouribility increased with an increase in RSAD and the value of intensified soil anti-scouribility (ΔAS) can be expressed by exponential equations, depending on the plant species. These equations were ΔAS=9.578 6 RSAD^0.8321 (R^2=0.951) for afforested Pinus tabulaeformis Carr.ΔAS=7.8087 RSAD^0.7894(R^2=0.974) for afforested Robinia pseudoacacia L., and ΔAS=9.256 6 RSAD^0.8707(R^2=0.899) for Bothriochloa ischemum L.     

Root distribution and morphology of maize seedlings as affected by tillage and fertilizer placement

Suboptimal soil conditions are known to result in poor early growth of maize (Zea mays L.) in no-tillage (NT) systems in contrast with conventional tillage (CT) systems. However, most studies have generally focused on maize roots at later growth stages and/or do not give details on root morphology. In a 2-year field study at two locations (silt loam and loam soils) in the Swiss midlands, we investigated the impacts of tillage intensity, NT vs. CT, and NP-fertilizer sidebanding on the morphology, vertical and horizontal distribution, and nutrient uptake of maize roots at the V6 growth stage. The length density (RLD) and the length per diameter-class distribution (LDD) of the roots were determined from soil cores taken to a depth of 0.5 m and at distances of 0.05 and 0.15 m from both sides of the maize row. The temperature of the topsoil was lower, and the bulk density and penetration resistance were greater in the topsoil of NT compared with CT. The growth and the development of the shoot were slower in NT. RLD was greater and the mean root diameter smaller in CT than in NT, while the vertical and horizontal distribution of roots did not differ between CT and NT. RLD increased in the zone enriched by the sidebanded fertilizer, independent of the tillage system, but LDD did not change. The poorer growth of the roots and shoots of maize seedlings was presumably caused by the lower topsoil temperature in NT rather than by mechanical impedance. The placement of a starter fertilizer at planting under NT is emphasized.     

Soil Anti-Scouribility Enhanced by Plant Roots

The magnitude of soil anti-scouribility depends on the physical condition of the soil. Plant roots can greatly enhance soil stability and anti-erodibility. A scouring experiment of undisturbed soil was conducted to investigate the effects of roots on soil anti-scouribility and its distribution in the soil profile. At the end of each erosion test, plant roots were collected from soil samples and root surface area was calculated by means of a computer image analysis system (CIAS). Root surface area density (RSAD), the surface area of the roots per unit of soil volume, was related to soil anti-scouribility. More than 83% of root surface area was concentrated in the 0 - 30 cm soil layer. Soil anti-scouribility increased with an increase in RSAD and the value of intensified soil anti-scouribility (△AS) can be expressed by exponential equations, depending on the plant species. These equations were △AS = 9.578 6 RSAD0.8321 (R2 = 0.951) for afforested Pinus tabulaeformis Cart., △AS = 7.808 7 RSAD0.7894 (R2 = 0.974) for afforested Robinia pseudoacacia L., and △AS = 9.256 6 RSAD0.8707 (R2 = 0.899) for Bothriochloa ischemum L.     

The distribution and abundance of wheat roots in a dense,structured subsoil – implications for water uptake

We analysed the abundance, spatial distribution and soil contact of wheat roots in dense, structured subsoil to determine whether incomplete extraction of subsoil water was due to root system limitations. Intact soil cores were collected to 1.6 m below wheat crops at maturity on a red Kandosol in southern Australia. Wheat roots, remnant roots, soil pores and root–soil contact were quantified at fresh breaks in the soil cores. In surface soil layers (<0.6 m) 30–40% of roots were clumped within pores and cracks in the soil, increasing to 85–100% in the subsoil (>0.6 m), where 44% of roots were in pores with at least three other roots. Most pores contained no roots, with occupancy declining from 20% in surface layers to 5% in subsoil. Wheat roots clumped into pores contacted the surrounding soil via numerous root hairs, whereas roots in cracks were appressed to the soil surface and had very few root hairs. Calculations assuming good root–soil contact indicated that root density was sufficient to extract available subsoil water, suggesting that uptake is constrained at the root–soil interface. To increase extraction of subsoil water, genetic targets could include increasing root–soil contact with denser root hairs, and increasing root proliferation to utilize existing soil pores.     

陇东旱塬苹果根系分布规律及生理特性对地表覆盖的响应

为探明陇东旱塬区不同覆盖物对苹果园土壤理化性状、根系分布及根系生理活性的影响,以14年生苹果树为试材,采用土壤剖面分层取样法,调查根系空间分布,并对根系生物量、根长、表面积等进行分析,测定根系活力、抗氧化酶类、活性氧代谢等相关生理指标,同时测定不同深度土层土壤容重、孔隙度等.结果表明: 覆草可有效增大土壤含水量、孔隙度、有机质含量,增幅分别为2.7%～11.6%、3.2%～27.7%、5.1%～36.0%,但土壤容重降低,为清耕(CK)的88.7%～96.4%.CK根系主要分布在距树干30～120 cm范围内的0～60 cm深土层中;覆草、覆膜处理主要分布在距树干0～150 cm、0～60 cm水平范围内的0～100 cm深土层中,以20～40 cm根系最为密集;覆膜处理细根总量仅为CK的96.4%,根系水平分布范围较CK有所减小,0～60 cm内细根占根系总量的51.6%.不同覆盖处理显著增强0～80 cm土层根系活力及抗氧化酶活性,其中覆草处理根系活力为CK的111.3%～136.7%.综合分析根系生长分布与生理活性、土壤理化性状等,认为覆草处理是陇东旱塬区苹果园较为适宜的地表覆盖方式.     

Citrus root responses to localized drying soil: A new approach to studying mycorrhizal effects on the roots of mature trees

Because fine roots tend to be concentrated at the soil surface, exposure to dry surface soil can have a large influence on patterns of root growth, death and respiration. We studied the effects of arbuscular mycorrhizas (AM) formation on specific root length (SRL), respiration and mortality of fine roots of bearing red grapefruit (Citrus paradisi Macf.) trees on Volkamer lemon (C. volkameriana Tan. & Pasq.) rootstock exposed to drying soil. For each tree, the fine roots were removed from two woody lateral roots, the roots were surface sterilized and then each woody root was placed in a separate pair of vertically divided and independently irrigated soil compartments. The two split-pot systems were filled with sterilized soil and one was inoculated with arbuscular mycorrhizal fungi (Glomus etunicatum/G. intraradices). New fine lateral roots that emerged from the woody laterals were permitted to grow inside the pots over a 10-month period. Irrigation was then removed from the top compartment for a 15-week period. At the end of the study, roots inoculated with AM fungi exhibited about 20% incidence of AM formation, whereas the uninoculated roots were completely void of AM fungi. Arbuscular mycorrhizal roots exhibited lower SRL, lower root/soil respiration and about 10% lower fine root mortality than nonmycorrhizal roots after 15 weeks of exposure to dry surface soil. This study demonstrates the feasibility of examining mycorrhizal effects on the fine roots of adult trees in the field using simple inexpensive methods.     

隔沟交替灌溉条件下玉米根系形态性状及结构分布

为揭示根系对土壤环境的适应机制,研究了隔沟交替灌溉条件下玉米根系形态性状及结构分布。以垄位和坡位的玉米根系为研究对象,利用Minirhizotrons法研究了根系（活/死根）的长度、直径、体积、表面积、根尖数和径级变化及其与土壤水分、土温和水分利用效率（WUE）的相关关系。结果表明,对于活根,在坡位非灌水区域复水后根系平均直径减小,而根系日均生长速率、单位面积土壤根系体积密度、根尖数和表面积均增大,并随灌水区域土壤水分的消退逐渐减小;对于死根,在坡位非灌水区域复水后根系日均死亡速率、根系体积密度、根尖数和表面积变化均减小,其中根系死亡速率和死根直径随土壤水分的消退逐渐降低,而死根体积密度、根尖数和表面积分布随土壤水分降低呈增大趋势;在垄位,根系形态分布趋势与坡位一致,除根系直径与与坡位比较接近外,其他根系形态值均小于坡位。将根系分成4个径级区间分析根系的形态特征,结果表明在根系长度和体积密度分布中以2.5－4.5 mm径级的根系所占比例最大,在根尖数和根系表面积分布中以0.0－2.5 mm径级的根系为主。通过显著性相关分析,死根直径、体积密度、活根表面积等根系形态与土壤含水率、土壤温度和WUE间均存在显著或极显著的正相关关系,部分根系形态指标（如根系的生长速率、活根体积密度）只与坡位土壤含水量、土壤温度具有明显的相关性,表明隔沟交替灌溉对坡位根系形态的调控作用比垄位显著。     

Use of a microtensiometer technique to study hydraulic lift in a sandy soil planted with pearl millet (Pennisetum americanum [L.] Leeke)

A pot experiment was carried out with pearl millet (Pennisetum americanum [L.] Leeke) growing in a sandy soil in which the upper (topsoil) and lower (subsoil) parts of the pots were separated by a perlite layer to prevent capillary water movement. Using microtensiometers a study was made to establish whether it was possible to measure hydraulic lift by which the upper part of the soil was rewetted when water was supplied exclusively to the lower part of the soil.Hydraulic lift occurred during the first seven days of the period of measurement, with a maximum water release to the soil of 2.7 Vol. % during one night (equivalent to 10.8 mL water in the top 10 cm of the soil profile). This magnitude was obtained at very high root length densities, so that water release from the roots would be expected to be much smaller under field conditions.Hydraulic lift ceased when the soil matric potential in the topsoil dropped below-10 kPa at the end of the light period and could not be re-established, neither by extending the dark period, nor after rewatering the topsoil. The disappearance of hydraulic lift could be explained in part through osmotic adaptation of plant roots and, thus prevention of water release from the roots in the topsoil. It is concluded that hydraulic lift may affect nutrient uptake from drying topsoil by extending the time period favourable for uptake from the topsoil.     

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.     

The role of soil hydraulic conductivity on the spatial and temporal variation of root water uptake in drip-irrigated corn

A multiplexed TDR system and a heat-pulse system for stem sap flow measurements were used to determine the spatial and temporal pattern of root water uptake in field-grown corn. The TDR probes, 0.15 and 0.30 m in length, were buried vertically in the soil profile to a depth of 0.95 m below the soil surface and heat-pulse sensors were installed on the plant base. Nocturnal readings from TDR probes were used successfully to differentiate the two components of moisture change: root uptake and net drainage. The instantaneous rate of water extraction by the plant measured by the heat-pulse system agreed well with the integrated rate of root water uptake measured frequently (at half-hour or hourly intervals) by the TDR probes. This agreement enabled further exploration into the cause of the evolution of the spatial and temporal patterns of root water uptake during a drying cycle. The results indicated that right after irrigation in the well-watered soil profile, it is the spatial distribution of the roots that mainly determines the typical pattern of root extraction, in addition to the fact that the roots near the plant base are more effective than those farther away. The higher density and effectiveness of the roots near the plant base dry the soil rapidly so that soil hydraulic conductivity soon becomes a limiting factor for water uptake. Further analysis revealed that a decrease in root uptake occurs near the plant base under a given atmospheric demand when the relative bulk soil hydraulic conductivity decreases to 0.002K _r. This suggests that low conductivity (high resistance) in the soil near the plant base is the initial cause for downward and lateral shifting of the root uptake pattern. Note that this critical value of hydraulic conductivity is not universal since it depends on the soil type and atmospheric water demand during the period under observation. Therefore, prior to the application of moisture content or suction head as measures of water availability or to control irrigation scheduling, it is suggested that these parameters be calibrated by the soil K() or K() curves, respectively, for the expected atmospheric water demand for the specific crop and growing period.     

土壤湿度对香樟幼苗根系生长和构型的影响

以1年生香樟(Cinnamomum camphora)幼苗为试材,设置对照组(CK)、中度干旱处理(M)、重度干旱处理(S)三个处理,比较不同土壤湿度下香樟幼苗不同时期地上部分生长和根系构型,探究香樟幼苗根系对不同土壤湿度的适应性及其耐旱机制。结果表明,中度和重度干旱处理组的香樟根系及地上部分干物质积累、根系长度、根系表面积、根系直径和根尖数均显著小于对照组(P<0.05)。同时干旱显著增加香樟幼苗的根系拓扑指数,降低香樟根的分形维数和平均分枝角度(P<0.05)。可见土壤湿度程度及处理时间显著影响香樟根系的生长及在土壤中的布局。较低土壤湿度可显著抑制根长的延长、根表面积扩大和根的增殖,且随着土壤湿度的继续降低以及处理时间延长,香樟幼苗根系的生长受到水分亏缺的抑制作用加重,根系建成成本增高的同时,根系分枝的复杂性降低,根系必须通过朝着更陡、更深的方向生长伸长来提高水分吸收效率。建议在园林绿化工程养护过程中制定科学的水分管理策略,以满足香樟生长过程中对土壤水分的需要。     

Understanding the impact of root morphology on overturning mechanisms: a modelling approach

BACKGROUND AND AIMS: The Finite Element Method (FEM) has been used in recent years to simulate overturning processes in trees. This study aimed at using FEM to determine the role of individual roots in tree anchorage with regard to different rooting patterns, and to estimate stress distribution in the soil and roots during overturning. METHODS: The FEM was used to carry out 2-D simulations of tree uprooting in saturated soft clay and loamy sand-like soil. The anchorage model consisted of a root system embedded in a soil block. Two root patterns were used and individual roots removed to determine their contribution to anchorage. KEY RESULTS: In clay-like soil the size of the root-soil plate formed during overturning was defined by the longest roots. Consequently, all other roots localized within this plate had no influence on anchorage strength. In sand-like soil, removing individual root elements altered anchorage resistance. This result was due to a modification of the shape and size of the root-soil plate, as well as the location of the rotation axis. The tap root and deeper roots had more influence on overturning resistance in sand-like soil compared with clay-like soil. Mechanical stresses were higher in the most superficial roots and also in leeward roots in sand-like soil. The relative difference in stresses between the upper and lower sides of lateral roots was sensitive to root insertion angle. Assuming that root eccentricity is a response to mechanical stresses, these results explain why eccentricity differs depending on root architecture. CONCLUSIONS: A simple 2-D Finite Element model was developed to better understand the mechanisms involved during tree overturning. It has been shown how root system morphology and soil mechanical properties can modify the shape of the root plate slip surface as well as the position of the rotation axis, which are major components of tree anchorage.     

典型黑土区主要树种根系构型特征及其对固土能力的影响

为量化典型黑土区主要树种根系构型特征,探究其对固土能力的影响,以该区分布较广的榆叶梅、小叶锦鸡儿、白桦、糖槭、红皮云杉、樟子松单株个体为研究对象,采用全根挖掘和WinRHIZO Pro LA2004分析系统相结合对其根系空间分布、几何形态、分形等特征进行测定,同时采用原位整株根系拉拔的方法量化根系垂直拉拔力。结果表明: 榆叶梅以倾斜根为主,小叶锦鸡儿、白桦、糖槭和红皮云杉以水平根为主,樟子松根系在水平和垂直分布上较为均衡;除白桦总根表面积和红皮云杉总根长外,灌木树种总根长、总根表面积显著大于乔木,落叶阔叶乔木总根长、总根表面积显著大于针叶常绿乔木,白桦总根体积显著大于小叶锦鸡儿、糖槭、红皮云杉和樟子松;榆叶梅、小叶锦鸡儿和白桦根系分形维数和分形丰度显著大于红皮云杉和樟子松;榆叶梅、小叶锦鸡儿和糖槭整株根系平均最大垂直拉拔力显著大于白桦、樟子松和红皮云杉。主要受根系总根长、总根表面积和倾斜根数量的影响,榆叶梅、小叶锦鸡儿和糖槭根系表现出较强的固土能力,可作为典型黑土区水土保持植被构建中优先选择的树种。     

The role of soil conditions in fine root ecomorphology in Norway spruce (Picea abies (L.) Karst.)

The present study is an attempt to investigate the pattern of morphological variability of the short roots of Norway spruce (Picea abies (L.) Karst.) growing in different soils. Five root parameters – diameter, length and dry weight of the root tip, root density (dry weight per water-saturated volume) and specific root area (absorbing area of dry weight unit) were studied with respect to 11 soil characteristics using CANOCO RDA analysis. The investigation was conducted in seven study areas in Estonia differing in site quality class and soil type. Ten root samples per study area were collected randomly from the forest floor and from the 20 cm soil surface layer. Eleven soil parameters were included in the study: humus content, specific soil surface area, field capacity, soil bulk density, pH (KCl and H₂O dilution's), N and Ca concentrations, Ca/Al and C/N ratios, and the decomposition rate of fine roots (<2 mm dia.). Root morphological characteristics most strongly related to the measured soil characteristics in the different sites were specific root area, root density and diameter of the short roots, the means varying from 29 to 42 m² kg⁻¹, from 310 to 540 kg m⁻³ and from 0.26 to 0.32 mm, respectively; root density being most sensitive. The most favourable site and soil types resulting in fine roots with morphological characteristics for optimizing nutrient uptake (e.g. low short root density and high specific root area) were Umbric Luvisol (Oxalis), Dystric Gleysol (Oxalis) and Gleyic Luvisol (Hepatica). These soil types correspond to highly productive natural forest stands of Norway spruce in Estonia. All measured soil variables explained 28% of total variance of the root characteristics. The most important variables related to root morphology were the humus content, field capacity and specific soil surface area. This revised version was published online in June 2006 with corrections to the Cover Date.     

The effects of lime and phosphorus on the function of wheat roots in acid top soils and subsoils

Two wheat varieties with differing aluminium tolerance were grown in pots of acid soil. Liming did not change significantly the amounts of chemically extractable P and K, but caused improved vegetative growth, increased inflow of P and K and reduced uptake of Al. Without lime, roots had a higher content and concentration of P than shoots; liming reversed this. Without lime the sensitive variety with a shorter root length had an Al inflow ten times that of the tolerant one: tolerance involves a mechanism for exlcuding Al. The inflow of P per unit inflow of Al (mol ratio) without lime was three times greater for the tolerant variety which therefore has more P to counteract the effects of Al. The same varieties were grown in two-layer soil columns, with a low P status and a limed topsoil and acid subsoil. Liming the subsoil improved plant growth but this was still restricted by low P availability. Addition of P to the topsoil caused good growth regardless of subsoil acidity: root growth increased in both layers and P (labelled with³²P) taken up from the topsoil was translocated to roots in the subsoil. This P inactivated root Al and allowed the roots to grow and take up more P from the acid subsoil with however a reduction in inflow. The sensitive variety was affected more by the acid subsoil and low P availability, had a similar ability to translocate P to subsoil roots but could not attain the growth rate of the tolerant wheat even with P and lime.     

Changes in root morphology of white lupin (Lupinus albus L.) and its adaptation to soils with heterogeneous alkaline/acid profiles

The ability of Lupinus albus L. to adapt to a heterogeneous soil profile containing acid subsoil below limed topsoil of the same type, and to utilize nutrients by significantly altering its root system structure, was investigated using specially constructed soil profile tubes. Plants grown in homogeneous acid profiles had the fastest growth while those grown in homogeneous limed-soil profiles showed the slowest growth and exhibited some chlorosis after 19 days. Limed topsoil combined with an acid subsoil profile initially retarded plant growth similar to that in a homogeneous limed soil. However, after 68 days significantly greater growth had occurred in the limed/acid soil treatment relative to the homogeneous limed soil, indicating plants had benefited from the acid subsoil stratum. Plants in the homogeneous limed soil profile had lower concentrations of P, Fe and Mn in shoots compared with those in heterogeneous soils. In contrast, the concentration of Ca increased by 74%, due mainly to an increase in the water-soluble Ca fraction. When grown in a heterogeneous limed/acid soil profile, concentrations of P, Ca, K, Mg, Fe, Mn and Zn in shoots were comparable to those grown in a soil with a homogeneous acid profile. Although total root production was lower in the homogeneous limed-soil profile compared to the acid-soil containing profiles, cluster root mass was maintained at a level comparable with that in acid soil. The roots in heterogeneous soil profiles exhibited extensive plasticity, demonstrating a root-type specific, morphological response to the soil conditions. Within the acid subsoil of a heterogeneous profile, there was a large increase in cluster root mass compared with non-cluster roots. The proliferation of cluster roots in acid soil below limed topsoil may enhance the plant's ability to exploit this soil and facilitate the cultivation of L. albus on limed soil. This revised version was published online in August 2006 with corrections to the Cover Date.