Root system architecture of Quercus pubescens trees growing on different sloping conditions

BACKGROUND AND AIMS: Plant roots' growth direction has important implications for plant development and survival; moreover it plays an effective and vital role in stabilizing weathered soil on a steep slope. The aim of this work was to assess the influence of slope on the architecture of woody root systems. METHODS: Five mature, single-stemmed Quercus pubescens trees growing on a steep slope and five on a shallow slope were excavated to a root diameter of 1 cm. A very precise numeric representation of the geometry and topology of structural root architecture was gained using a low-magnetic-field digitizing device (Fastrak, Polhemus). Several characteristics of root architecture were extracted by macros, including root volume, diameter, length, number, spatial position and branching order. KEY RESULTS: The diameter at breast height (dbh) was the best predictor of the root volume but had no correlation with length and number of roots. The slope affected the root volume for each branching order, and the basal cross-sectional area (CSA), number and length of the first-order roots. Number and length of the second- and third-order laterals were closely related in both conditions, although this relationship was closer in the shallow trees, suggesting the influence of a genetic control. Sloping trees showed a clustering tendency of the first- and second-order lateral roots in the up-slope direction, suggesting that the laterals rather than the taproots provide much of the anchorage. In a steep-slope condition, the taproot tapering was positively correlated with the asymmetry magnitude of first-order roots, indicating compensation between taproot and main lateral roots' clustering tendency. CONCLUSIONS: These results suggest that on a slope, on clayey soils, root asymmetry appears to be a consequence of several environmental factors such as inclination, shallow-slides and soil compactness. In addition, this adaptive growth seems to counteract the turning moment induced by the self-loading forces acting in slope conditions, and as a consequence improves the tree stability.     

Deep Phenotyping of Coarse Root Architecture in R. pseudoacacia Reveals That Tree Root System Plasticity Is Confined within Its Architectural Model

This study aims at assessing the influence of slope angle and multi-directional flexing and their interaction on the root architecture of Robinia pseudoacacia seedlings, with a particular focus on architectural model and trait plasticity. 36 trees were grown from seed in containers inclined at 0° (control) or 45° (slope) in a glasshouse. The shoots of half the plants were gently flexed for 5 minutes a day. After 6 months, root systems were excavated and digitized in 3D, and biomass measured. Over 100 root architectural traits were determined. Both slope and flexing increased significantly plant size. Non-flexed trees on 45° slopes developed shallow roots which were largely aligned perpendicular to the slope. Compared to the controls, flexed trees on 0° slopes possessed a shorter and thicker taproot held in place by regularly distributed long and thin lateral roots. Flexed trees on the 45° slope also developed a thick vertically aligned taproot, with more volume allocated to upslope surface lateral roots, due to the greater soil volume uphill. We show that there is an inherent root system architectural model, but that a certain number of traits are highly plastic. This plasticity will permit root architectural design to be modified depending on external mechanical signals perceived by young trees.     

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.     

Control of soil physical properties and response ofBrassica rapa L. seedling roots

A system was designed, constructed, tested, and used to growBrassica rapa L. seedling roots which were exposed to O₂ concentrations from 0 to 0.21 mol mol⁻¹, water potentials from 0 to −80 kPa, temperatures from 10 to 34°C, and mechanical impedance from 0 to 20.8 kPa. The experimental design was a central composite rotatable design with seven replications of the center point. Measurements were taken of taproot length, taproot diameter at the point of initiation of root hairs (diameter 1), and one cm above the first measurement (diameter 2), and total length and number of first-order laterals. Temperature had the greatest effect on seedling root growth, with linear and quadratic temperature effects significant for all root measurements except taproot diameter 2 which just had a significant linear effect. Water potential had a significant linear effect on lateral length and number of laterals and a significant quadratic effect on taproot diameter 1. Mechanical impedance had a significant effect only on taproot diameter 2. Oxygen was not significant for any root measurement. The mechanical impedance by water potential interaction was significant for taproot length and taproot diameter 1. A temperature optimum was found for taproot length, taproot diameter 1, lateral length, and lateral number, at 26.0, 42.5, 26.5, and 26.4°C, respectively. Taproot diameter 1 had a water potential optimum at −36.5 kPa, whereas taproot diameter 2 had a mechanical impedance optimum at 12.5 kPa. A growth cell designed for this study allows independent control of soil strength, water potential, oxygen concentration, and temperature. Thus, the cell provides the capability which was demonstrated forBrassica rapa L. to grow seedling roots under complete control of the soil physical properties.     

Gravitropic response growth angle and vertical distribution of roots of wheat (Triticum aestivum L.)

Recent work on root distribution, growth angles and gravitropic responses in Japanese cultivars of winter wheat are reviewed. Vertical distribution of roots, which influences the environmental stress tolerance of plants, was observed in the 12 Japanese cultivars in the field. The root depth index (RDI: the depth at which 50% of the root length has been reached) differed among the cultivars at the stem elongation stage. Since the RDI was closely related to the growth angle of seminal roots obtained in a pot experiment, it was assumed that growth angle is useful for predicting vertical root distribution among wheat genotypes. Gravitropic responses of the primary seminal root of 133 Japanese wheat cultivars assessed by measuring the growth angle in agar medium, were larger in the northern Japanese cultivars and smaller in the southern ones. It was also found that the geographical variation resulted from the wheat breeding process, i.e. genotypes with limited gravitropic responses of roots had been selected in the southern part of Japan where excessive soil moisture is one of the most serious problems.     

Sesame hairy root cultures for extra-cellular production of a recombinant fungal phytase

Sesame (Sesamum indicum L.) hairy roots were transformed with a fungal (Aspergillus) phytase and their culture conditions were surveyed for the extra-cellular production of the recombinant phytase protein in shake flasks. Kanamycin resistance of sesame hairy roots was observed at 50 μg ml⁻¹ kanamycin sulfate and southern hybridization analysis confirmed the existence of the phytase gene in the hairy root genomic DNA. The continuous dark condition was more effective for both the root growth and phytase production than light. Slightly higher root growth was determined at 30 °C than 26 °C in Murashige & Skoog (MS) medium supplemented with 3% sucrose, while the final phytase production was greatest in MS medium with 5 or 3% sucrose at both temperatures of 26 and at 30 °C. Among the culture media used, full-strength MS medium was exclusively efficient for production of the recombinant phytase. Most rapid increase rates in both the root growth and phytase production were detected at the 4th week of the culture periods and thereafter their rates began to decrease. Our results indicated that 5–6-week culture periods may be necessary for the maximal phytase production. Western analysis revealed that even though the phytase proteins expressed were measured with greater activities in the liquid medium than in the root tissues, they were still retained in the tissues.     

Unravelling the response of poplar (Populus nigra) roots to mechanical stress imposed by bending

Abstract

Mechanical stress is a widespread environmental condition that can be caused by several factors (i.e. gravity, touch, wind, soil density, soil compaction and grazing, slope) and that can severely affect plant stability. In response to mechanical stress and to improve their anchorage, plants have developed complex mechanisms to detect mechanical perturbation and to induce a suite of modifications at anatomical, physiological, biochemical, biophysical and molecular level. Although it is well recognized that one of the primary functions of root systems is to anchor the plant to the soil, root response to mechanical stresses have been investigated mainly at morphological and biomechanical level, whereas investigations about the molecular mechanisms underlying these important alterations are still in an initial stage. We have used an experimental system in which the taproot poplar seedlings are bent to simulate mechanical perturbation to begin investigate the mechanisms involved in root response to mechanical stress. The results reported herein show that, in response to bending, the poplar root changes its morphology by emitting new lateral roots, and its biomechanical properties by increasing the root biomass and lignin synthesis. In addition, using a proteomic approach, we found that several proteins involved in the signal transduction pathway, detoxification and metabolism are up-regulated and/or down-regulated in the bent root. These results provide new insight into the obscure field of woody root response to mechanical stress, and can serve as a basis for future investigations aimed at unravelling the complex mechanism involved in the reaction of root biology to environmental stress.     

The proteome of Populus nigra woody root: response to bending

Background and Aims

Morphological and biomechanical alterations occurring in woody roots of many plant species in response to mechanical stresses are well documented; however, little is known about the molecular mechanisms regulating these important alterations. The first forest tree genome to be decoded is that of Populus, thereby providing a tool with which to investigate the mechanisms controlling adaptation of woody roots to changing environments. The aim of this study was to use a proteomic approach to investigate the response of Populus nigra woody taproot to mechanical stress.

Methods

To simulate mechanical perturbations, the taproots of 30 one-year-old seedlings were bent to an angle of 90 ° using a steel net. A spatial and temporal two-dimensional proteome map of the taproot axis was obtained. We compared the events occurring in the above-bending, central bending and below-bending sectors of the taproot.

Key Results

The first poplar woody taproot proteome map is reported here; a total of 207 proteins were identified. Spatial and temporal proteomic analysis revealed that factors involved in plant defence, metabolism, reaction wood formation and lateral root development were differentially expressed in the various sectors of bent vs. control roots, seemingly in relation to the distribution of mechanical forces along the stressed woody taproots. A complex interplay among different signal transduction pathways involving reactive oxygen species appears to modulate these responses.

Conclusions

Poplar woody root uses different temporal and spatial mechanisms to respond to mechanical stress. Long-term bending treatment seem to reinforce the defence machinery, thereby enabling the taproot to better overcome winter and to be ready to resume growth earlier than controls.     

Willow water uptake and shoot extension growth in response to nutrient and moisture on a clay landfill cap soil

Extension growth of willow (Salix viminalis L.) and changes in soil water were measured in lysimeters containing clay and sandy loam soils with different amendment and watering treatments. No water uptake was found below 0.3 m in the nutritionally poor unamended clay; amendment with organic matter to 0.4 m depth resulted in water extraction down to 0.5 m depth whereas in the sandy loam, there was greater extraction from all depths down to 0.6 m. With water stress, wilting of plants occurred when the volumetric soil water content at 0.1 m was about 31% in the clay and 22% in the sandy loam. Compared with shoots on plants in the amended clay, those in the unamended treatment showed reduced extension growth, little increase in stem basal area (SBA) and a small shoot leaf area, resulting from a reduced number of leaves shoot⁻¹ and a small average area leaf⁻¹. Water stress also reduced shoot extension growth, SBA gain and the leaf area on extension growth. Shoot growth rates were significantly correlated with air temperature and base temperatures between 2.0 and 7.6 °C were indicated for the different treatments. These studies have helped to explain some of the large treatment effects described previously on biomass production and plant leaf area.     

川滇高山栎灌丛萌生过程中的营养元素供应动态

萌生更新是森林更新的重要方式, 是硬叶栎林受到干扰后植被恢复的主要机制。以位于青藏高原东南缘的川西折多山东坡川滇高山栎(Quercus aquifoliodes)灌丛为研究对象, 调查分析了砍伐后灌丛萌生过程中基株根系和萌株生物量动态、营养元素含量, 以及基株根系和土壤对萌株生长过程中的营养元素供应动态。结果表明, 川滇高山栎灌丛平均地上和地下生物量分别为(11.25 ± 0.92) t·hm^-2和(34.85 ± 2.02) t·hm^-2, 具有较大的根冠比(3.10:1); 萌生过程中, 萌株生物量呈线性增加趋势, 以灌丛活细根生物量变化为最大, 其次是活中根和活粗根, 树桩和根蔸生物量变化最小; 萌生过程中, 灌丛细根和中根N、P含量表现为先增加、后降低的变化趋势, 萌生初期树桩、粗根和根蔸中N和K的含量明显下降, 根蔸中Ca含量略有下降, 而P没有明显下降, 根系Mg含量变化幅度较大, 灌丛地下根系储存了较多的营养元素; 土壤、树桩、粗根和根蔸是川滇高山栎灌丛砍伐后0-120天萌生生长的主要营养来源, 砍伐后60天, 萌株生长所需的营养除K元素主要来源于根系外, 其余营养元素主要来源于土壤; 在砍伐后60-120天, 基株根系对萌株生长所需的N、K和Ca贡献较大, 而对P和Mg的贡献较小; 在砍伐后120-180天, 根系除K元素对萌生生长还保持较大的贡献外, 对其余营养元素的贡献均较小。高山栎林管理要注重加强地下根系的保护。     

Direct heat injury of roots of woody swamp species

1. 1.Three month-old seedlings of Taxodium distichum, Nyssa aquatica, Cephalanthus occidentalis and cuttings of Salix nigra were acclimated to simulated natural swamp conditions (ambient temperature, saturated soil) and then used to assess direct high temperature injury of root tissue.

2. 2.Electrolyte leakage from excised root tissue exposed for 30 min to temperatures ranging from 30 to 66°C was used to assess cellular injury.

3. 3.The relationship between leakage and temperature was sigmoidal for each species.

4. 4.Inflection point temperatures on the response curves, ranged from 45.4 to 51.0°C, were species-specific, and indicated differences in thermal tolerance of root membranes.

5. 5.Root of C. occidentalis and N. aquatica were more heat tolerant than roots of T. distichum and S. nigra.

Plagiogravitropism of maize roots

The direction of root growth can be studied by analyzing the trajectories of roots growing in soil. Both the primary seminal root and nodal roots of maize attain a preferred, or liminal, angle of growth that deviates from the vertical. These roots are said to be plagiogravitropic. Experiments using plants grown in soil-filled boxes revealed that the primary seminal root is truly plagiogravitropic. It shows both positive and negative gravitropism in response to gravity stimuli and tends to maintain its direction even after growing around obstacles. These are experimental results suggesting that plagiogravitropic growth is controlled by internal factors. The orientation of the grain affects the establishment of the liminal angle of the primary seminal root, and both the position of their node of origin and the root diameter are closely related to the plagiogravitropic behaviour of nodal roots. Several external factors are also known to influence plagiogravitropism. Low soil water content causes a decrease in the angle of growth and soil mechanical resistance suppresses the gravitropic curvature. Plagiogravitropic behaviour of both seminal and nodal roots plays a significant role in shaping the root system.     

表施和混施污泥对团花根系生长的影响

林地利用被认为是污泥资源化利用的重要方式,但施用污泥对林木根系生长的影响报道较少。本研究通过根箱试验,分析表施和混施10%污泥对速生树种团花不同土层根系形态、土壤pH值和电导率动态变化及根系重金属含量的影响,并拟合土壤pH值、电导率和根系重金属含量与根长的关系。结果表明: 与不施污泥(对照)相比,混施污泥显著抑制了团花根长、根表面积和根体积增长,混施污泥120和240 d后,0～20 cm土层总根长分别为不施污泥的76.9%和67.4%;表施污泥对团花根长和根表面积的影响不显著,但显著提高了根体积。混施污泥显著提高了土壤pH值和电导率及根系重金属含量,混施污泥0～20和20～40 cm土层根系镉含量分别是不施污泥的11.5和10.0倍。线性回归拟合分析显示,不同处理0～20 cm土层的电导率与根长均呈显著负相关;表施和混施污泥根系镉含量与根长呈极显著负相关。上述结果表明,混施污泥抑制了团花根系生长,这可能是由于混施污泥提高了土壤电导率和根系镉含量所致,而表施污泥对团花根系生长的作用不明显。     

Changes in plasmalemma K+Mg2+ -ATPase dephosphorylating activity and H+ transport in relation to seasonal growth and freezing tolerance of Festuca pratensis Huds

Changes in plasmalemma K⁺Mg²⁺-ATPase dephosphorylating activity and H⁺ transport were examined in freezing-tolerant and non-tolerant genotypes of the perennial grass species Festuca pratensis Huds. Enzyme activity and ΔμH⁺ were measured in plasmalemma fractions isolated from basal nodes and roots. Three types of experiments were undertaken: (i) a field experiment, utilizing the seasonal growth and cessation cycle of a perennial plant; (ii) a cold acclimation experiment in hydroponics; and (iii) an instant freezing test. A specific fluctuation in K⁺Mg²⁺-ATPase activity was found throughout the seasonal growth of the plants (i). The K⁺Mg²⁺-ATPase activity peaks for both the basal node and the root plasmalemma were determined early in the spring before the renewal of growth. The lowest activity values in roots occurred at the time approaching flowering, and in basal nodes at the transition into the growth cessation. The K⁺Mg²⁺-ATPase activity was approximately 50% lower in the basal node plasmalemma of freezing-tolerant plants than of non-tolerant ones, when assessed at the optimal growth stage in hydroponics. In hydroponics (ii) and in the freezing test (iii), temperature stress was followed by a more pronounced change in the level of K⁺Mg²⁺-ATPase activity than in that of H⁺ transport, and this change was more clearly differentiated in the basal node plasmalemma of contrasting genotypes than in the roots. Stress response was manifested differently in freezing-tolerant and non-tolerant plants at cold acclimation (4–2 °C) and at freezing (−8 °C) temperatures. Proton transport regulation via coupled changes in the hydrolysed ATP/transported proton ratio, as an attribute of freezing-tolerant plants, is discussed.     

Effect of cyclic wetting and drying of a soil on root hair growth of maize roots

The objective of this study was to investigate the effect of cyclic soil wetting and drying on maize (Zea mays L.) root hair growth. Three soils, Chalmers silty clay loam (Typic Haplaquolls), Raub silt loam (Aquic Argiudolls) and Aubbeenaubbee sandy loam (Aric Ochraqualfs) and two soil moisture contents, −175 (M₀) and −7.5 kPa (M₁), were used to study root hair growth in a controlled-climate chamber. Increasing soil moisture after 7d from M₀ and M₁ resulted in a cessation of root hair growth behind the root cap while drying the soil after 7d from M₁ and M₀ promoted root hair growth on new but not old or existing roots. By maintaining liquid continuity under cyclic wetting and drying of a soil, root hairs may be of far greater significance to the nutrition of the plant than originally thought. Journal Paper No. 11023, Purdue Univ. Agric. Exp. Stn., W. Lafayette, IN 47907. Contribution from the Dep. of Agron.     

The influence of calcium and pH on growth in primary roots of Zea mays

Hasenstein, K. H. and Evans, M. L. 1988. The influence of calcium and pH on growth in primary roots of Zea mays. - Physiol. Plant. 72: 466–470.
We investigated the interaction of Ca²⁺ and pH on root elongation in Zea mays L. cv. B73 × Missouri 17 and cv. Merit. Seedlings were raised to contain high levels of Ca²⁺ (HC, imbibed and raised in 10 m M CaCl₂) or low levels of Ca²⁺ (LC, imbibed and raised in distilled water). In HC roots, lowering the pH (5 m M MES/Tris) from 6.5 to 4.5 resulted in strong, long-lasting growth promotion. Surprisingly, increasing the pH from 6.5 to 8.5 also resulted in strong growth promotion. In LC roots acidification of the medium (pH 6.5 to 4.5) resulted in transient growth stimulation followed by a gradual decline in the growth rate toward zero. Exposure of LC roots to high pH (pH shift from 6.5 to 8.5) also promoted growth. Addition of EGTA resulted in strong growth promotion in both LC and HC roots. The ability of EGTA to stimulate growth appeared not to be related to H⁺ release from EGTA upon Ca²⁺ chelation since, 1) LC roots showed a strong and prolonged response to EGTA, but only a transient response to acid pH, and 2) promotion of growth by EGTA was observed in strongly buffered solutions. We also examined the pH dependence of the release of ⁴⁵Ca²⁺ from roots of 3-day-old seedlings grown from grains imbibed in ⁴⁵Ca²⁺. Release of ⁴⁵Ca²⁺ from the root into agar blocks placed on the root surface was greater the more acidic the pH of the blocks. The results indicate that Ca²⁺ may be necessary for the acid growth response in roots.     

Modification of root architecture in woody plants is possible for the presence of two different mechanisms of lateral root production: The effect of slope in Spartium junceum L. seedlings

Abstract

This paper investigates the modification of root architecture of Spartium junceum L. seedlings grown in slope condition. It is reported that 50% of the total number of lateral roots are concentrated in few centimetres of the taproot near the collar. The anatomical analysis of transverse sections along the taproot axis reveals that this taproot zone is characterised by two types of lateral roots: one with a trace extending to the centre of the vascular cylinder by following the path of a medullar ray; one with a trace which ends in the vascular cambium. The first type may be lateral roots originated from the taproot primary structure; the second type seems to be lateral roots developing later when a secondary structure has completely substituted the primary structure. The emission of this second type of lateral roots seems to be strongly controlled by environmental conditions with considerable consequences upon the overall root architecture. In the example reported in this paper, young plants growing under mechanical stress due to a slope develop asymmetric root architecture with lateral roots elongating in two prevalent directions: up-slope or down-slope. This asymmetric architecture is produced in the zone of the taproot where a secondary structure is present and represents the plant response to the need of increasing its anchorage strength.     

刈割对紫花苜蓿根系生长影响的初步分析

研究了不同刈割次数对紫花苜蓿根系发育能力的影响。结果表明：刈割抑制紫花苜蓿主根的纵向(垂直)生长,而促进其横向(直径)生长,并使紫花苜蓿侧根发生总数减少和垂直分布的总体格局向表层集中,随着刈割次数的增加,侧根集中分布在土壤表层的比例也增加;刈割使根系生物量和体积总量减少,并随着刈割次数的增加,这种减少趋势更加明显,同时在土壤中的垂直分布变浅。     

A study of the tensile force required to pull wheat roots from soil

Experiments were carried out to determine the tensile properties of mature wheat roots and the force necessary to pull roots from undisturbed soils at a range of moisture contents using an Instron materials testing rig. Roots decreased in cross sectional area from 1.5 to 0.1 mm²and in tensile strength from 7.0 to 2.3 Newtons (N) along the first 12 cm of their length. Breaking strain was constant along the root but the breaking stress increased. Increased seeding rate decreased root diameter and tensile strength but plant growth regulators and fertiliser nitrogen level did not affect the tensile properties. Roots were pulled from mesh tubes of soil (25 mm ± 75 mm) into which they had grown. The peak loads for sandy clay loam and sandy loam soils were 4.8 and 3.9 N respectively and increasing the soil moisture from 17% to 26% reduced the peak load from 5.2 N to 3.5 N. With the drier soil the inner stele pulled free leaving the outer periderm in the soil in a higher proportion of the roots indicating a stronger root/soil bond than in the wetter soil. The load us displacement relationship when pulling roots from soil resembled that for a fibre reinforced composite material. The presence of branch roots resulted in an uneven trace in which there were a number of sub-peaks as branch roots gave way. It is suggested that soil wetting could contribute to lodging of wheat by reducing the resistance of roots to slippage and breaking.     

Effect of phosphorus deficiency on growth angle of basal roots in Phaseolus vulgaris

Root architectural plasticity might be an important factor in the acquisition by plants of immobile nutrients such as phosphorus (P). In this study, we examined the effect of P availability on the orientation of basal roots with respect to gravity, and thereby on the growth angle of these roots of common bean (Phaseolus vulgaris L.). In one set of studies the growth angle of basal roots of bean seedlings was measured over time. Sixteen bean genotypes were examined; six showed a decrease in root orientation with respect to gravity in low P media, one increased orientation, and nine showed no difference within 5 d of basal root emergence. Bean taproots also showed decreased root orientation with respect to gravity in low P. Growth angle after 5 d was correlated with field performance of contrasting genotypes in low P tropical soils. Mineral deficiencies other than P did not cause changes in root angle. In a split pouch system that provided high or low P solution to different parts of the root system, the decrease in root angle in low P was found to be a response to global P availability, and not local to the portion of the root system in low P. Effects of P availability on root angle were associated with reduced shoot P concentration, but preceded effects on plant biomass accumulation and leaf area expansion. Results from growth pouches for genotype G 19833 were confirmed using a solid-phase buffered sand-culture system supplying P at three levels. Pea (Pisum sativum), soybean (Glycine max Williams), chickpea (Cicer arietinum), lima bean (Phaseolus lunatus), and lentil (Lens culinaris) were grown with and without P; soybean and pea also showed decreased basal root angles in low P.