Transient three-dimensional modeling of soil water and solute transport with simultaneous root growth,root water and nutrient uptake

A three-dimensional solute transport model was developed and linked to a three-dimensional transient model for soil water flow and root growth. The simulation domain is discretized into a grid of finite elements by which the soil physical properties are spatially distributed. Solute transport modeling includes passive and active nutrient uptake by roots as well as zero- and first-order source/sink terms. Root water uptake modeling accounts for matric and osmotic potential effects on water and passive nutrient uptake. Root age effects on root water and nutrient uptake activity have been included, as well as the influence of nutrient deficiency and ion toxicity on root growth. Examples illustrate simulations with different levels of model complexity, depending on the amount of information available to the user. At the simplest level, root growth is simulated as a function of mechanical soil strength only. Application of the intermediate level with root water and nutrient uptake simulates the influence of timing and amount of NO3 application on leaching. The most comprehensive level includes simulation of root and shoot growth as influenced by soil water and nutrient status, temperature, and dynamic allocation of assimilate to root and shoot.     

Variation in phosphorus uptake efficiency by genotypes of cowpea (Vigna unguiculata) due to differences in root and root hair length and induced rhizosphere processes

The lengths of roots and root hairs and the extent of root-induced processes affect phosphorus (P) uptake efficiency by plants. To assess the influence of variation in the lengths of roots and root hairs and rhizosphere processes on the efficiency of soil phosphorus (P) uptake, a pot experiment with a low-P soil and eight selected genotypes of cowpea (Vigna unguiculata (L) WALP) was conducted. Root length, root diameter and root hair length were measured to estimate the soil volume exploited by roots and root hairs. The total soil P was considered as a pool of Olsen-P, extractable with 0.5 M NaHCO₃ at pH 8.5, and a pool of non-Olsen-P. Model calculations were made to estimate P uptake originated from Olsen-P in the root hair zone and the Olsen-P moving by diffusion into the root hair cylinder and non-Olsen-P uptake. The mean uptake rate of P and the mean rate of non-Olsen-P depletion were also estimated. The genotypes differed significantly in lengths of roots and root hairs, and in P uptake, P uptake rates and growth. From 6 to 85% of total P uptake in the soil volume exploited by roots and root hairs was absorbed from the pool of non-Olsen-P. This indicates a considerable activity of root-induced rhizosphere processes. Hence the large differences show that traits for more P uptake-efficient plants exist in the tested cowpea genotypes. This opens the possibility to breed for more P uptake-efficient varieties as a way to bring more sparingly soluble soil P into cycling in crop production and obtain capitalisation of soil P reserves.     

Regulation of root hair density by phosphorus availability in Arabidopsis thaliana

We characterized the response of root hair density to phosphorus (P) availability in Arabidopsis thaliana. Arabidopsis plants were grown aseptically in growth media with varied phosphorus concentrations, ranging from 1 mmol m⁻³ to 2000 mmol m⁻³ phosphorus. Root hair density (number of root hairs per mm of root length) was analysed starting at 7 d of growth. Root hair density was highly regulated by phosphorus availability, increasing significantly in roots exposed to low-phosphorus availability. The initial root hairs produced by the radicle were not sensitive to phosphorus availability, but began to respond after 9 d of growth. Root hair density was about five times greater in low phosphorus (1 mmol m⁻³) than in high phosphorus (1000 mmol m⁻³) media. Root hair density decreased logarithmically in response to increasing phosphorus concentrations within that range. Root hair density also increased in response to deficiencies of several other nutrients, but not as strongly as to low phosphorus. Indoleacetic acid (IAA), the auxin transport inhibitor 2-(p-chlorophenoxy)-2-methylpropionic acid (CMPA), the ethylene precursor 1-aminocyclopropane-1-carboxylic acid (ACC), and the ethylene synthesis inhibitor amino-oxyacetic acid (AOA) all increased root hair density under high phosphorus but had very little effect under low phosphorus. Low phosphorus significantly changed root anatomy, causing a 9% increase in root diameter, a 31% decrease in the cross-sectional area of individual trichoblasts, a 40% decrease in the cross-sectional area of individual atrichoblasts, and 45% more cortical cells in cross-section. The larger number of cortical cells and smaller epidermal cell size in low phosphorus roots increased the number of trichoblast files from eight to 12. Two-thirds of increased root hair density in low phosphorus roots was caused by increased likelihood of trichoblasts to form hairs, and 33% of the increase was accounted for by changes in low phosphorus root anatomy resulting in an increased number of trichoblast files. These results show that phosphorus availability can fundamentally alter root anatomy, leading to changes in root hair density, which are presumably important for phosphorus acquisition.     

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.     

Influences of root distribution and growth on predicted water uptake and interspecific competition

Summary Root distribution and growth measured in the field were incorporated into a water uptake model for the CAM succulent Agave deserti and its nurse plant Hilaria rigida, a common desert bunchgrass. Agave deserti responds to the infrequent rainfalls of the Sonoran Desert by extending its existing established roots and by producing new roots. Most of such root growth was completed within one month after soil rewetting, total root length of A. deserti increasing by 84% for a seedling and by 58% for a mediumsized plant in the summer. Root growth in the winter with its lower soil temperatures was approximately half as much as in the summer. For a 15-year period, predicted annual root growth of A. deserti varied more than 18-fold because of annual variations in rainfall amount and pattern as well as seasonal variation in soil temperature. Predicted annual water uptake varied 47-fold over the same period. The nurse plant, which is crucial for establishment of A. deserti seedlings, reduced seedling water uptake by 38% during an average rainfall year. Lowering the location of the root system of a medium-sized A. deserti by 0.24 m reduced its simulated annual water uptake by about 25%, reflecting the importance of shallow roots for this desert succulent. Lowering the root system of a medium-sized H. rigida by 0.28 m increased the simulated annual water uptake of an associated A. deserti seedling by 17%, further indicating the influence of root overlap on competition for water.     

The root sap of birch on a podzol

Summary Root sap analyses were made on birch trees growing in a strongly podzolised heath soil. In the early summer the concentration of calcium and phosphorus appears to vary with the quantity of sap produced, and is independent of root environment. Later in the summer, however, this relationship no longer holds; there now being a connection between rooting zone and nutrient concentration. Roots reaching theC horizon absorb greater quantities of bases, whilst the superficial roots show higher concentrations of phosphorus and ammoniacal nitrogen. Throughout the season potassium uptake remains practically independent of both sap yield and rooting zone. In these soils young pine trees do not penetrate to the subsoil, but a comparison of sap analyses from surface roots of pine and birch shows that birch sap is richer in most nutrients. This, together with the ability of the birch to exploit the subsoil, may be a contributory factor to the soil improving properties of a birch crop.     

Membrane-mediated decrease in root exudation responsible for phorphorus inhibition of vesicular-arbuscular mycorrhiza formation

The mechanism responsible for phosphorus inhibition of vesicular-arbuscular mycorrhiza formation in sudangrass (Sorghum vulgare Pers.) was investigated in a phosphorus-deficient sandy soil (0.5 micrograms phosphorus per gram soil) amended with increasing levels of phosphorus as superphosphate (0, 28, 56, 228 micrograms per gram soil). The root phosphorus content of 4-week-old plants was correlated with the amount of phosphorus added to the soil. Root exudation of amino acids and reducing sugars was greater for plants grown in phosphorus-deficient soil than for those grown in the phosphorus-treated soils. The increase in exudation corresponded with changes in membrane permeability of phosphorus-deficient roots, as measured by K⁺ (⁸⁶Rb) efflux, rather than with changes in root content of reducing sugars and amino acids. The roots of phosphorus-deficient plants inoculated at 4 weeks with Glomus fasciculatus were 88% infected after 9 weeks as compared to less than 25% infection in phosphorus-sufficient roots; these differences were correlated with root exudation at the time of inoculation. For plants grown in phosphorus-deficient soil, infection by vesicular-arbuscular mycorrhizae increased root phosphorus which resulted in a decrease in root membrane permeability and exudation compared to nonmycorrhizal plants. It is proposed that, under low phosphorus nutrition, increased root membrane permeability leads to net loss of metabolites at sufficient levels to sustain the germination and growth of the mycorrhizal fungus during pre- and postinfection. Subsequently, mycorrhizal infection leads to improvement of root phosphorus nutrition and a reduction in membrane-mediated loss of root metabolites.     

亚热带典型树种根毛特征及其与共生真菌的关系

根毛和共生真菌增加了吸收面积,提高了植物获取磷等土壤资源的能力。由于野外原位观测根表微观结构较为困难,吸收细根、根毛、共生真菌如何相互作用并适应土壤资源供应,缺乏相应的数据和理论。该研究以受磷限制的亚热带森林为对象,选取了21种典型树种,定量了根毛存在情况、属性变异,分析了根毛形态特征与共生真菌侵染率、吸收细根功能属性之间的关系,探讨了根表结构对低磷土壤的响应和适应格局。结果表明:1)在亚热带森林根毛不是普遍存在的, 21个树种中仅发现7个树种存有根毛, 4个为丛枝菌根(AM)树种, 3个为外生菌根(ECM)树种。其中,马尾松(Pinus massoniana)根毛出现率最高,为86%;2)菌根类型是理解根-根毛-共生真菌关系的关键,AM树种根毛密度与共生真菌侵染率正相关,但ECM树种根毛直径与共生真菌侵染率负相关; 3) AM树种根毛长度和根毛直径、ECM树种根毛出现率与土壤有效磷含量呈负相关关系。该研究揭示了不同菌根类型树种根毛-共生真菌-根属性的格局及相互作用,为精细理解养分获取策略奠定了基础。     

Genotypic differences in the presence of hairs on roots and gynophores of peanuts (Arachis hypogaea L.) and their significance for phosphorus uptake

Root hairs substantially increase the surface area of plant roots with positive effects for phosphorus (P) uptake, but the ability of peanuts to form root hairs has been questioned. The aim was to examine hair development on roots and gynophores of a variety of peanut genotypes and to relate genotypic differences in hair formation to differences in P uptake. Five out of eighteen genotypes completely lacked hairs on both organs whereas others consistently developed hairs on roots and gynophores, although with considerable variation in hair density. The ability to form root hairs as well as root hair density concurred with the presence and density of hairs on gynophores, suggesting a possible connection between both developmental processes. The contribution of root hairs to P uptake was studied in three genotypes differing in hair density. The final amount of P taken up by roots did not differ between genotypes but two distinct P uptake strategies could be identified. The genotype lacking root hairs maintained P uptake due to the development of a large root system whereas densely covered roots of genotype 'Wasedairyu' were three times as efficient in extracting P from a P-deficient soil. Furthermore P uptake through gynophores contributed about 20% to the total P uptake of Wasedairyu but only insignificant amounts to other genotypes. The ability to form hairs on roots and gynophores can therefore be seen as an adaptation to low P availability and if combined with a large root system, could substantially increase the tolerance of peanuts to P deficiency.     

Effect of phosphorus availability on basal root shallowness in common bean

Root gravitropism may be an important element of plant response to phosphorus availability because it determines root foraging in fertile topsoil horizons, and thereby phosphorus acquisition. In this study we seek to test this hypothesis in both two dimensional paper growth pouch and three-dimensional solid media of sand and soil cultures. Five common bean (Phaseolus vulgaris L.) genotypes with contrasting adaptation to low phosphorus availability were evaluated in growth pouches over 6 days of growth, and in sand culture and soil culture over 4 weeks of growth. In all three media, phosphorus availability regulated the gravitropic response of basal roots in a genotype-dependent manner. In pouches, sand, and soil, the phosphorus-inefficient genotype DOR 364 had deeper roots with phosphorus stress, whereas the phosphorus-efficient genotype G19833 responded to phosphorus stress by producing shallower roots. Genotypes were most responsive to phosphorus stress in sand culture, where relative root allocation to the 0–3- and 3–6-cm horizons increased 50% with phosphorus stress, and varied 300% (3–6 cm) to 500% (0–3 cm) among genotypes. Our results indicate that (1) phosphorus availability regulates root gravitropic growth in both paper and solid media, (2) responses observed in young seedlings continue throughout vegetative growth, (3) the response of root gravitropism to phosphorus availability varies among genotypes, and (4) genotypic adaptation to low phosphorus availability is correlated with the ability to allocate roots to shallow soil horizons under phosphorus stress.     

The efficiency of Arabidopsis thaliana (Brassicaceae) root hairs in phosphorus acquisition

Arabidopsis thaliana root hairs grow longer and denser in response to low-phosphorus availability. In addition, plants with the root hair response acquire more phosphorus than mutants that have root hairs that do not respond to phosphorus limiting conditions. The purpose of this experiment was to determine the efficiency of root hairs in phosphorus acquisition at high- and low-phosphorus availability. Root hair growth, root growth, root respiration, plant phosphorus uptake, and plant phosphorus content of 3-wk-old wild-type Arabidopsis (WS) were compared to two root hair mutants (rhd6 and rhd2) under high (54 mmol/m) and low (0.4 mmol/m) phosphorus availability. A cost-benefit analysis was constructed from the measurements to determine root hair efficiency. Under high-phosphorus availability, root hairs did not have an effect on any of the parameters measured. Under low-phosphorus availability, wild-type Arabidopsis had greater total root surface area, shoot biomass, phosphorus per root length, and specific phosphorus uptake. The cost-benefit analysis shows that under low phosphorus, wild-type roots acquire more phosphorus for every unit of carbon respired or unit of phosphorus invested into the roots than the mutants. We conclude that the response of root hairs to low-phosphorus availability is an efficient strategy for phosphorus acquisition.     

Influence of root density on the critical soil water potential

Estimation of root water uptake in crops is important for making many other agricultural predictions. This estimation often involves two assumptions: (1) that a critical soil water potential exists which is constant for a given combination of soil and crop and which does not depend on root length density, and (2) that the local root water uptake at given soil water potential is proportional to root length density. Recent results of both mathematical modeling and computer tomography show that these assumptions may not be valid when the soil water potential is averaged over a volume of soil containing roots. We tested these assumptions for plants with distinctly different root systems. Root water uptake rates and the critical soil water potential values were determined in several adjacent soil layers for horse bean (Vicia faba) and oat (Avena sativa) grown in lysimeters, and for field-grown cotton (Gossypium L.), maize (Zea mays) and alfalfa (Medicago sativa L.) crops. Root water uptake was calculated from the water balance of each layer in lysimeters. Water uptake rate was proportional to root length density at high soil water potentials, for both horse bean and oat plants, but root water uptake did not depend on root density for horse bean at potentials lower than −25 kPa. We observed a linear dependency of a critical soil water potential on the logarithm of root length density for all plants studied. Soil texture modified the critical water potential values, but not the linearity of the relationship. B E Clothier Section editor     

Localized supply of phosphorus induces root morphological and architectural changes of rice in split and stratified soil cultures

A localized supply of phosphorus may affect root morphology and architecture, and thereby affect phosphorus uptake by rice plants. In the present study, we attempted to test this hypothesis using two rice cultivars representing upland and lowland ecotypes grown in specially designed split and stratified soil cultures with a low-phosphorus red soil. Our data indicate that a localized supply of phosphorus increased both total root length and root fineness, particularly in the high-phosphorus zone. In split culture, plants roots tended to preferentially grow on the high-phosphorus zone, with about 70–75% of the total root length allocated to the high-phosphorus compartment. The total root length on the high-phosphorus side in the split-phosphorus treatment was significantly longer than that in the homogenously high-phosphorus treatment, implying that a phosphorus-deficiency signal from the low-phosphorus side may stimulate the growth of the roots located in the high-phosphorus zone. In stratified soil culture, changes in root morphology and architecture were also observed as indicated by increased total root length, root fineness and relative root allocation in the high-phosphorus layers, again suggesting altered root morphology and preferential root proliferation in the high-phosphorus regions. The induced changes in root morphology and architecture by localized phosphorus supply may have both physiological significance and practical implications in that plants can meet the demand for phosphorus with parts of the roots reaching the high-phosphorus zone, hence localized fertilization methods such as side dressing or banded application of phosphorus fertilizers may both minimize phosphorus fixation by the soil and increase phosphorus uptake efficiency from the fertilizers.     

Root cap removal increases root penetration resistance in maize (Zea mays L)

The root cap assists the passage of the root through soil by means of its slimy mucilage secretion and by the sloughing of its outer cells. The root penetration resistance of decapped primary roots of maize (Zea mays L. cv. Mephisto) was compared with that of intact roots in loose (dry bulk density 1.0 g cm-3; penetration resistance 0.06 MPa) and compact soil (1.4 g cm-3; penetration resistance 1.0 MPa), to evaluate the contribution of the cap to decreasing the impedance to root growth. Root elongation rate and diameter were the same for decapped and intact roots when the plants were grown in loose soil. In compacted soil, however, the elongation rate of decapped roots was only about half that of intact roots, whilst the diameter was 30% larger. Root penetration resistances of intact and decapped seminal axis were 0.31 and 0.52 MPa, respectively, when the roots were grown in compacted soil. These results indicated that the presence of a root cap alleviates much of the mechanical impedance to root penetration, and enables roots to grow faster in compacted soils.     

Reduced root cortical burden improves growth and grain yield under low phosphorus availability in maize

Root phenes and phene states that reduce the metabolic cost of soil exploration may improve plant growth under low phosphorus availability. We tested the hypothesis that under low phosphorus, reduced living cortical area (LCA) would increase soil exploration, phosphorus capture, biomass, and grain yield. Maize genotypes contrasting in LCA were grown in the field and in greenhouse mesocosms under optimal and suboptimal phosphorus regimes. Percent LCA in nodal roots ranged from 25% to 67%. Plants with 0.2 mm² less LCA under low phosphorus had 75% less root segment respiration, 54% less root phosphorus content, rooted 20 cm deeper, allocated up to four times more roots between 60 and 120 cm depth, had between 20% and 150% more biomass, 35–40% greater leaf phosphorus content, and 60% greater grain yield compared with plants with high LCA. Low‐LCA plants had up to 55% less arbuscular mycorrhizal colonization in axial roots, but this decrease was not correlated with biomass or phosphorus content. The LCA components cortical cell file number and cortical cell size were important for biomass and phosphorus content under low phosphorus. These results are consistent with the hypothesis that root phenes that decrease the metabolic cost of soil exploration are adaptive under phosphorus stress.     

小麦开花期灌水对土壤养分及根系分布的影响

本研究通过分析开花期灌水对小麦产量、植株养分分配和土壤养分分布的影响及其与根系特性的关系,为小麦充分利用水肥资源提供理论支撑。以抗旱高产品种‘洛麦28'和高光效品种‘百农207'为材料,采用2 m深土柱栽培方法,设置开花期灌水(T₁)和开花期不灌水(T₂)两个水分处理,测定了不同组织器官、不同土层土壤氮、磷、钾含量及根系分布特性等指标。结果表明: 小麦收获期土壤中铵态氮、速效磷和速效钾主要分布在0～80 cm土层中,硝态氮主要分布在80 cm以下土层中,开花期灌水促进小麦吸收0～60 cm土层的铵态氮、速效磷、速效钾和80 cm以下土层的硝态氮,减少了硝态氮向深层土壤的淋溶;小麦根系主要集中在0～60 cm土层中,随土壤深度的增加而减少。成熟期干物质积累量、全氮和全磷主要分配在小麦籽粒中,而全钾主要分配在茎秆中;开花期灌水显著增加了小麦百粒重,提高了小麦产量;根系形态指标与土壤硝态氮在0～40 cm土层中呈显著负相关,与土壤铵态氮在80～100 cm土层中呈极显著正相关,与土壤速效磷在0～100 cm土层中呈显著正相关。开花期灌水促进了根系在小麦生育末期对土壤养分的充分吸收,延长了养分从营养器官向生殖器官的转运功能期,使营养器官中的养分充分地转运到籽粒中去,增加小麦粒重,进而提高产量。     

Root-shoot signalling in sunflower plants with confined root systems

Sunflower plants were grown hydroponically under controlled conditions with the root systems confined in small containers. Root confinement inhibited the growth of sunflower plants as indicated by reduction in both leaf and cotyledon area and root and shoot fresh weight. This effect was more pronounced in shoots. Root confinement favored the accumulation of potassium in the roots and shoots, and the exudation of potassium and water in excised roots. Xylem sap from root confined plants inhibited cotyledon expansion as revealed by bioassay with decapited sunflower seedlings. In addition decapited control plants incubated in ABA solution also showed cotyledon growth reduction. Xylem sap ABA analysis indicated a 7-times higher concentration in root confined than control plants. Our results suggest the synthesis of a chemical signal in the roots of plants subjected to mechanical stress which can be responsible for the inhibition of plant growth.     

A split-pot experiment with sorghum to test a root water uptake partitioning model

Correct modeling of root water uptake partitioning over depth is an important issue in hydrological and crop growth models. Recently a physically based model to describe root water uptake was developed at single root scale and upscaled to the root system scale considering a homogeneous distribution of roots per soil layer. Root water uptake partitioning is calculated over soil layers or compartments as a function of respective soil hydraulic conditions, specifically the soil matric flux potential, root characteristics and a root system efficiency factor to compensate for within-layer root system heterogeneities. The performance of this model was tested in an experiment performed in two-compartment split-pot lysimeters with sorghum plants. The compartments were submitted to different irrigation cycles resulting in contrasting water contents over time. The root system efficiency factor was determined to be about 0.05. Release of water from roots to soil was predicted and observed on several occasions during the experiment; however, model predictions suggested root water release to occur more often and at a higher rate than observed. This may be due to not considering internal root system resistances, thus overestimating the ease with which roots can act as conductors of water. Excluding these erroneous predictions from the dataset, statistical indices show model performance to be of good quality.     

Influence of temperature and water potential on root growth of white oak

Root growth of white oak ( Quercus alba L.) was observed under field conditions using a rhizotron. The effects of temperature, soil water potential, and leaf water potential were evaluated on three measures of root growth and development: root elongation rate, number of growing roots, and root growth intensity (sum of projected root area compared to the total root viewing area). Root elongation rate was linearly related to changes in soil temperature and soil water potential. At soil temperatures less than 17deg;C, temperature was the dominant factor affecting rate of growth, bat at temperatures greater than 17°C soil water potential became the important factor. Unlike root elongation rate, the number of growing roots and root growth intensity increased at cold soil temperatures (8°C) and at soil water potentials of-0.3 to -0.8 MPa. At high soil water potentials (-0.1 MPa) root elongation rate reached a maximum while the number of growing roots and root growth intensity were low. These differences showed that root growth and development were not exclusively affected by the soil environment. In addition, the relationship between root growth and predawn leaf water potential suggested that root growth was a contributing factor to the drought resistance of white oak.     

Effects of soil structural discontinuity on root and shoot growth and water use of maize

The role of roots penetrating various undisturbed soil horizons beneath loose layer in water use and shoot growth of maize was evaluated in greenhouse experiment. 18 undisturbed soil columns 20 cm in diameter and 20 cm in height were taken from the depths 30–50 cm and 50–70 cm from a Brown Lowland soil, a Pseudogley and a Brown Andosol (3 columns from each depth and soil). Initial resistance to penetration in undisturbed soil horizons varied from 2.5 to 8.9 MPa while that in the loose layer was 0.01 MPa. The undisturbed horizons had a major effect on vertical arrangement of roots. Root length density in loose layer varied from 96 to 126 km m^-3 while in adjacent stronger top layers of undisturbed horizons from 1.6 to 20.0 km m^-3 with higher values in upper horizons of each soil. For specific root length, the corresponding ranges were 79.4–107.7 m g^-1 (on dry basis) and 38.2–63.7 m g^-1, respectively. Ratios of root dry weight per unit volume of soil between loose and adjacent undisturbed layers were much lower than those of root length density indicating that roots in undisturbed horizons were produced with considerably higher partition of assimilates. Root size in undisturbed horizons relative to total roots was from 1.1 to 38.1% while water use from the horizons was from 54.1 to 74.0%. Total water use and shoot growth were positively correlated with root length in undisturbed soil horizons. There was no correlation between shoot growth and water use from the loose layers.