Shading and the capture of localized soil nutrients: nutrient contents,carbohydrates, and root uptake kinetics of a perennial tussock grass

Summary The ability to exploit spatial and temporal heterogeneity in soil resources can be one factor important to the competitive balance of plants. Competition above-ground may limit selective plant responses to below-ground heterogeneity, since mechanisms such as root proliferation and alterations in uptake kinetics are energy-dependent processes. We studied the effect of shading on the ability of the perennial tussock grassAgropyron desertorum to take up nutrients from enriched soil microsites in two consecutive growing seasons. Roots of unshaded plants selectively increased phosphate uptake capacity in enriched soil microsites (mean increases of up to 73%), but shading eliminated this response. There were no changes in ammonium uptake capacity for roots in control and enriched patches for either shaded or unshaded plants. The 9-day shade treatments significantly reduced total nonstructural carbohydrate (TNC) concentrations for roots in 1990, but had no apparent effect on root carbohydrates in 1991 despite dramatic reductions in shoot TNC and fructan concentrations. Enrichment of the soil patches resulted in significantly greater phosphate concentrations in roots of both shaded and unshaded plants, with less dramatic differences for nitrogen and no changes in potassium concentrations. In many respects the shaded plants did surprisingly well, at least in terms of apparent nutrient acquisition. The effects of aboveground competition on nutrient demand, energy requirements, and belowground processes are discussed for plants exploiting soil resource heterogeneity.     

Effect of soil moisture and phosphate level on root hair growth of corn roots

Summary Root hairs have been shown to enhance P uptake by plants growing in low P soil. Little is known of the factors controlling root hair growth. The objective of this study was to investigate the influence of soil moisture and P level on root hair growth of corn (Zea mays L.). The effect of volumetric soil moistures of 22% (M₀), 27% (M₁), and 32% (M₂) and soil (Raub silt loam, Aquic Argiudoll) P levels of, 0.81 (P₀), 12.1 (P₁), 21.6 (P₂), 48.7 (P₃), and 203.3 (P₄) mol P L^–1 initially in the soil solution, on shoot and root growth, P uptake, and root hair growth of corn was studied in a series of pot experiments in a controlled climate chamber. Root hair growth was affected more by soil moisture than soil P. The percentage of total root length with root hairs and the density and length of root hairs on the root sections having root hairs all increased as soil moisture was reduced from M₂ to M₀. No relationship was found between root hair length and soil P. Density of root hairs, however, was found to decrease with an increase in soil P. No correlation was found between root hair growth parameters and plant P content, further suggesting P plays a secondary role to moisture in regulating root hair growth in soils. The increase in root hair growth appears to be a response by the plant to stress as yield and P uptake by corn grown at M₀ were only 0.47 to 0.82, and 0.34 to 0.74, respectively, of that measured at M₁ across the five soil P levels. The increase in root hair growth at M₀, which represents an increase of 2.76 to 4.03 in root surface area, could offset, in part, the reduced rate of root growth, which was the primary reason for reduced P uptake under limited soil moisture conditions.Journal Paper No. 10,066 Purdue Univ. Agric. Exp. Stn., W. Lafayette, IN 47907. Contribution from the Dep. of Agron. This paper was supported in part by a grant from the Tennessee Valley Authority.     

Root proliferation characteristics of seven perennial arid-land grasses in nutrient-enriched microsites

We compared root proliferation in fertilized microsites among seven cultivars of five commonly planted cool-desert perennial grass species that differ in productivity and competitive ability. In a greenhouse experiment on nutrient-limited plants, one soil microsite in each pot received distilled water (control) and a second microsite received a rich, complete nutrient solution (fertilized). Roots in and adjacent to the microsites were mapped on Mylar windows for 22 days after the injections to determine the magnitude and timing of response in root length relative growth rates (RGRs). Because we provided adequate water, used a high level of fertilization in the treatment microsites, and conducted the experiments during rapid vegetative growth, the results provide a measure of the relative capacities and maximal rates of the grasses responses to enriched microsites. Root samples were harvested from control and fertilized microsites at the end of the experiment to determine the morphological basis of the proliferation response. In all seven grasses fine roots proliferated in the fertilized microsites faster than in the control microsites. The grasses did not differ in the timing of their response which showed a peak 7–8 days after injection. Although one species, Pseudoroegneria spicata cv. Goldar, had higher maximum root length RGR and higher RGR ratio (RGR in fertilized to RGR in control microsites) 7–8 days after injection, the seven grasses did not differ significantly in the magnitude of root length RGR response to fertilizer integrated over the 22 day experiment. The grasses also did not differ significantly in root morphological changes in fertilized mocrosites. Compared to roots in control microsites, roots in fertilized microsites had greater specific root length, length of secondary roots per length of main axis, number of lateral and sublateral roots per length of main axis, and mean lateral root length. Root proliferation was mainly the result of increased lateral branching and lateral root growth in all seven grasses. The consistency of root proliferation responses among these seven cultivars suggests that differences in the capacity for, maximum rate, or morphological basis of root proliferation are not directly related to ecological characteristics such as productivity and competitive ability. Other aspects of root response to nutrient enrichment, such as differential responses as a function of microsite nutrient concentration, plant phenology, plant nutrient status, or specific nutrient element(s), may still be important, but further experiments are required to determine whether different responses to enriched soil microsites among species correspond with know species differences in ecological characteristics.     

Effects of liming and mycorrhizal colonization on soil phosphate depletion and phosphate uptake by maize (Zea mays L.) and soybean (Glycine max L.) grown in two tropical acid soils

The effects of liming and inoculation with the arbuscular mycorrhizal fungus, Glomus intraradices Schenck and Smith on the uptake of phosphate (P) by maize (Zea mays L.) and soybean (Glycine max [L.] Merr.) and on depletion of inorganic phosphate fractions in rhizosphere soil (Al-P, Fe-P, and Ca-P) were studied in flat plastic containers using two acid soils, an Oxisol and an Ultisol, from Indonesia. The bulk soil pH was adjusted in both soils to 4.7, 5.6, and 6.4 by liming with different amounts of CaCO₃.In both soils, liming increased shoot dry weight, total root length, and mycorrhizal colonization of roots in the two plant species. Mycorrhizal inoculation significantly increased root dry weight in some cases, but much more markedly increased shoot dry weight and P concentration in shoot and roots, and also the calculated P uptake per unit root length. In the rhizosphere soil of mycorrhizal and non-mycorrhizal plants, the depletion of Al-P, Fe-P, and Ca-P depended in some cases on the soil pH. At all pH levels, the extent of P depletion in the rhizosphere soil was greater in mycorrhizal than in non-mycorrhizal plants. Despite these quantitative differences in exploitation of soil P, mycorrhizal roots used the same inorganic P sources as non-mycorrhizal roots. These results do not suggest that mycorrhizal roots have specific properties for P solubilization. Rather, the efficient P uptake from soil solution by the roots determines the effectiveness of the use of the different soil P sources. The results indicate also that both liming and mycorrhizal colonization are important for enhancing P uptake and plant growth in tropical acid soils.     

Studies on in vitro selection of Fe-efficient lines in sugarcane

The objective of this research was to show how a mechanistic uptake model that accurately predicts phosphorus (P) uptake by maize (Zea mays L.) in a pot experiment may be used to evaluate the reasons for the differences in P availability observed when soil pH is varied. The model predicts P uptake by integrating soil P supply by mass flow and diffusion; size, shape and growth rate of roots; and P uptake kinetics of the root. The P supply parameters of the model that may be affected by soil pH are P_li, initial P concentration in the soil solution; b, the buffer power of P in the soil, P_si, for P_li, and D_e, effective diffusion coefficient. The effect of these changes on P uptake was predicted with the model by using measured values of the three soil supply parameters and of size, shape, and growth rate of roots and keeping the other parameters at values characteristic of maize. Values for three soil supply parameters can be calculated from measurements of P_li, P_si, and , volumetric water content. The predictions of the model closely agreed with observed uptake when form of P present at the higher pH's was accounted for. There was a significant positive correlation (r=0.94) between P_li and observed P uptake and a significant negative correlation (-0.93) between P_si and observed P uptake. The use of the model demonstrated the significance of P form and the importance of P_li in P uptake. It also showed importance of root growth rate.     

Effects of plant species on phosphorus availability in a range of grassland soils

Vegetative conversion from grass to forest may influence soil nutrient dynamics and availability. A short-term (40 weeks) glasshouse experiment was carried out to investigate the impacts of ryegrass (Lolium perenne) and radiata pine (Pinus radiata) on soil phosphorus (P) availability in 15 grassland soils collected across New Zealand using ³³P isotopic exchange kinetics (IEK) and chemical extraction methods. Results from this study showed that radiata pine took up more P (4.5–33.5 mg P pot^–1) than ryegrass (1.1–15.6 mg pot^–1) from the soil except in the Temuka soil in which the level of available P (e.g., E _1min P_i, bicarbonate extractable P_i) was very high. Radiata pine tended to be better able to access different forms of soil P, compared with ryegrass. There were no significant differences in the level of water soluble P (Cp, intensity factor) between soils under ryegrass and radiata pine, but the levels of Cp were generally lower compared with original soils due to plant uptake. The growth of both ryegrass and radiata pine resulted in the redistribution of soil P from the slowly exchangeable P_i pool (E _{> 10m} P_i, reduced by 31.8% on the average) to the rapidly exchangeable P_i (E _1min-1d P_i, E _1d-10m P_i) pools in most soils. The values of R/r ₁ (the capacity factor) were also generally greater in most soils under radiata pine compared with ryegrass. Specific P mineralisation rates were significantly greater for soils under radiata pine (8.4–21.9%) compared with ryegrass (0.5–10.8%), indicating that the growth of radiata pine enhanced mineralisation of soil organic P. This may partly be ascribed to greater root phosphatase activity for radiata pine than for ryegrass. Plant species × soil type interactions for most soil variables measured indicate that the impacts of plant species on soil P dynamics was strongly influenced by soil properties.     

Cluster-root formation, carboxylate exudation and proton release of Lupinus pilosus Murr. as affected by medium pH and P deficiency

The study examined the interactive effect of pH and P supply on cluster-root formation, carboxylate exudation and proton release by an alkaline-tolerant lupin species (Lupinus pilosus Murr.) in nutrient solution. The plants were exposed to 1 (P₁, deficient) and 50 μM P (P₅₀, adequate) for 34 days in nutrient solution at either pH 5.6 or 7.8. Plant biomass was not influenced by pH at P₁, but at P₅₀ shoot and root dry weights were 23 and 18% higher, respectively, at pH 7.8 than at pH 5.6. There was no significant difference in plant biomass between two P treatments regardless of medium pH. Phosphorus deficiency increased significantly the number of the second-order lateral roots compared with the P₅₀ treatment. Both total root length and specific root length of plants grown at pH 5.6 were higher than those at pH 7.8 regardless of P supply. Cluster roots were formed at P₁, but cluster-root number was 2-fold higher at pH 7.8 than pH 5.6. Roots released 16 and 31% more protons at pH 5.6 and 7.8, respectively, in P₁ than in P₅₀ treatments, and the rate of proton release followed the similar pattern. At pH 5.6, citrate exudation rate was 0.39 μmol g⁻¹ root DW h⁻¹ at P₁, but was under the detection limit at P₅₀; at pH 7.8, it was 2.4-fold higher in P₁ than in P₅₀ plants. High pH significantly increased citrate exudation rate in comparison to pH 5.6. The uptake of anions P and S was inhibited at P₁ and high pH increased cations Na, Mg and Ca uptake. The results suggested that enhanced cluster-root formation, proton release and citrate exudation may account for the mechanism of efficient P acquisition by alkaline-tolerant L. pilosus well adapted to calcareous soils. Cluster-root formation and citrate exudation in L. pilosus can be altered by medium pH and P deficiency. Phosphorus deficiency-induced proton release may be associated with the reduced anion uptake, but high pH-induced proton release may be partly attributed to increased cation uptake.     

The timing and degree of root proliferation in fertile-soil microsites for three cold-desert perennials

Summary Root proliferation in nutrient-rich soil patches is an important mechanism facilitating nutrient capture by plants. Although the phenomenon of root proliferation is well documented, the specific timing of this proliferation has not been investigated. We studied the timing and degree of root proliferation for three perennial species common to the Great Basin region of North America: a shrub, Artemisia tridentata, a native tussock grass, Agropyron spicatum, and an introduced tussock grass, Agropyron desertorum. One day after we applied nutrient solution to small soil patches, the mean relative growth rate of Agropyron desertorum roots in these soil patches was two to four times greater than for roots of the same plants in soil patches reated with distilled water. Most of the increased root growth came from thin, laterally branching roots within the patches. This rapid and striking root proliferation by Agropyron desertorum occurred in response to N-P-K enrichment as well as to P or N enrichment alone. A less competitive bunchgrass, Agrophyron spicatum, showed no tendency to proliferate roots in enriched soil patches during these two-week experiments. The shrub Artemisia tridentata proliferated roots within one day of initial solution injection in the N-enrichment experiment, but root proliferation of this species was more gradual and less consistent in the N-P-K and P-enrichment experiments, respectively. The ability of Agropyron desertorum to proliferate roots rapidly may partly explain both its general competitive success and its superior ability to exploit soil nutrients compared to Agropyron spicatum in Great Basin rangelands of North America.     

The effect of phosphate rock dissolution on soil chemical properties and wheat seedling root elongation

Soils of the Appalachian region of the United States are acidic and deficient in P. North Carolina phosphate rock (PR), a highly substituted fluoroapatite, should be quite reactive in these soils, allowing it to serve both as a source of P and a potential ameliorant of soil acidity. An experiment was conducted to evaluate the influence of PR dissolution on soil chemical properties and wheat (Triticum aestivum cv. Hart) seedling root elongation. Ten treatments including nine rates of PR (0, 12.5, 25, 50, 100, 200, 400, 800, and 1600 mg P kg^-1) and a CaCO₃ (1000 mg kg^-1) control were mixed with two acidic soils, moistened to a level corresponding to 33 kPa moisture tension and incubated for 30 days. Pregerminated wheat seedlings were grown for three days in the PR treated soils and the CaCO₃ control. Root length was significantly (P<0.05) increased both by PR treatments and CaCO₃, indicating that PR dissolution was ameliorating soil acidity. The PR treatments increased soil pH, exchangeable Ca, and soil solution Ca while lowering exchangeable Al and 0.01 M CaCl₂ extractable soil Al. Root growth in PR treatments was best described by an exponential equation (P<0.01) containing 0.01 M CaCl₂ extractable Al. The PR dissolution did not reduce total soil solution Al, but did release Al complexing anions into soil solution, which along with increased pH, shifted Al speciation from toxic to nontoxic forms. These results suggest that North Carolina PR should contribute to amelioration of soil acidity in acidic, low CEC soils of the Appalachian region.     

Architectural Tradeoffs between Adventitious and Basal Roots for Phosphorus Acquisition

Adventitious rooting contributes to efficient phosphorus acquisition by enhancing topsoil foraging. However, metabolic investment in adventitious roots may retard the development of other root classes such as basal roots, which are also important for phosphorus acquisition. In this study we quantitatively assessed the potential effects of adventitious rooting on basal root growth and whole plant phosphorus acquisition in young bean plants. The geometric simulation model SimRoot was used to dynamically model root systems with varying architecture and C availability growing for 21 days at 3 planting depths in 3 soil types with contrasting nutrient mobility. Simulated root architectures, tradeoffs between adventitious and basal root growth, and phosphorus acquisition were validated with empirical measurements. Phosphorus acquisition and phosphorus acquisition efficiency (defined as mol phosphorus acquired per mol C allocated to roots) were estimated for plants growing in soil in which phosphorus availability was uniform with depth or was greatest in the topsoil, as occurs in most natural soils. Phosphorus acquisition and acquisition efficiency increased with increasing allocation to adventitious roots in stratified soil, due to increased phosphorus depletion of surface soil. In uniform soil, increased adventitious rooting decreased phosphorus acquisition by reducing the growth of lateral roots arising from the tap root and basal roots. The benefit of adventitious roots for phosphorus acquisition was dependent on the specific respiration rate of adventitious roots as well as on whether overall C allocation to root growth was increased, as occurs in plants under phosphorus stress, or was lower, as observed in unstressed plants. In stratified soil, adventitious rooting reduced the growth of tap and basal lateral roots, yet phosphorus acquisition increased by up to 10% when total C allocation to roots was high and adventitious root respiration was similar to that in basal roots. With C allocation to roots decreased by 38%, adventitious roots still increased phosphorus acquisition by 5%. Allocation to adventitious roots enhanced phosphorus acquisition and efficiency as long as the specific respiration of adventitious roots was similar to that of basal roots and less than twice that of tap roots. When adventitious roots were assigned greater specific respiration rates, increased adventitious rooting reduced phosphorus acquisition and efficiency by diverting carbohydrate from other root types. Varying the phosphorus diffusion coefficient to reflect varying mobilities in different soil types had little effect on the value of adventitious rooting for phosphorus acquisition. Adventitious roots benefited plants regardless of basal root growth angle. Seed planting depth only affected phosphorus uptake and efficiency when seed was planted below the high phosphorus surface stratum. Our results confirm the importance of root respiration in nutrient foraging strategies, and demonstrate functional tradeoffs among distinct components of the root system. These results will be useful in developing ideotypes for more nutrient efficient crops.     

Calcium and temperature effects on seedling exudation and root rot infection of common bean on an acid sandy soil

Soil born fungi such as Phytium ultimum, Fusarium ssp., and Rhizoctonia solani (Kühn) severely restrict stand establishment of common bean (Phaseolus vulgaris L.) on acid soils of the Tropics. Calcium application is known to alleviate fungal infection in many legumes but the causes are still unclear. To investigate environmental factors and physiological mechanisms involved, growth chamber experiments were conducted with an acid sandy soil from Mexico. Treatments were soil liming at a rate of 0.67 g Ca(OH)₂ kg^-1, gypsum application at 0.49 g CaSO₄ 2H₂O kg^-1 soil placed around the seed, and an untreated control. Beans were grown under three temperature regimes with constant night and one constant day vs. two sinusoidal day temperatures. To examine patterns of seed and seedling exudation at regular intervals leachates of germinating seeds were collected on filter paper soaked with equilibrium solutions from soils of the three treatments. The severity of root rot in the control treatment was highest when plants were stressed by temperature extremes. At a sinusoidal day temperature peaking at 40°C soil liming and gypsum application to the seed increased the number of healthy seedlings similarly by over 60%. However, only liming which effectively eliminated growth constraints by low pH and high aluminum concentrations led to an increase in hypocotyl elongation by 22% and in total root length by 8%. Both calcium amendments increased the calcium and potassium contents in the hypocotyl tissue. From seeds exposed to the equilibrium solution of unlimed soil with pH 3.7, 1 mM Ca, and 0.6 mM Al considerable amounts of amino acids and carbohydrates were leached. In contrast, exposure to the equilibrium solution from limed soil with pH 4.3, 3 mM Ca, and negligible concentrations of Al led to a net uptake of amino acids and decreased leaching of carbohydrates. Exposure to the equilibrium solution of the gypsum treatment with pH 3.6, 20 mM Ca, and 1.2 mM Al resulted in a somewhat smaller net uptake of amino acids compared to liming. During germination pH around the seeds steeply increased in the untreated control but significantly less with both amendments. The results indicate that pH and the Ca/Al ratio in the soil solution around bean seeds determine their pattern of exudation and solute uptake. For bean germination and early growth on acid soils locally placed application of small amounts of gypsum as seed pelleting seems as effective as soil liming in reducing the incidence of root rot. The results indicate that this may be accomplished by decreasing the amount of leachates available for fungal development.     

Chemistry of Cr in some Swedish soils

The effect of Si(OH)₄ on Cr toxicity and elemental concentrations in ryegrass were investigated in a growth chamber using an acid and a neutral mineral soil. Each soil was treated with 50 mg Cr, as CrO₃, kg⁻¹ soil dry weight, singly, or in combination with 25 mg Si as Si(OH)₄. Plants growing in unamended soils were used as controls. Chromium toxicity, expressed as decrease in shoot or root dry weight, was increased by the Si. This increase was accompanied by a higher Cr uptake particularly on the acid soil. The shoot and root dry weights were significantly correlated (P=1%) with the concentration of Cr, where r=−0.80 and −0.65, respectively. Uptake of Al, Cu, Fe, P and Zn did not show any consistent relationship to the magnitude of Cr toxicity.     

Soil acidity and nutrient deficiency in central Amazonian heath forest soils

Experiments were carried out to test the effects of liming and nutrient additions on plant growth and soil processes such as C and N mineralisation in three contrasting forest types in central Amazonia: the stunted facies of heath forest (SHF), the tall facies of heath forest (THF) and the surrounding lowland evergreen rain forest (LERF). Calcium-carbonate additions increased soil respiration in the field plots in the SHF; in laboratory incubations, soil respiration was higher in the SHF when soils were fertilised with N, and in THF and LERF after S additions. The addition of N alone or in different combinations generally induced a net immobilisation of soil N. Net nitrification increased during the incubation in SHF and THF soils fertilised with N+P, and in LERF soils fertilised with either N, or P, or CaCO₃. In a field experiment using ingrowth bags, a higher fine root production was observed in all forest types when bags were fertilised with CaCl₂ or CaCO₃, suggesting that Ca may be a limiting nutrient in these soils. Calcium-carbonate addition in a glasshouse bioassay experiment with rice showed an overall positive effect on the survival and growth of the seedlings. In other treatments where soil pH was not raised, the rice showed acute toxicity symptoms, poor root and shoot growth and high mortality. Similar results were yielded in a field experiment, using naturally established seedlings in the field plots in SHF, THF and LERF. It is concluded that the acute H⁺ ion toxicity is a major growth-limiting factor for non-adapted plants in heath forest soils in central Amazonia.     

Growth promotion of maize and lettuce by phosphate-solubilizing Rhizobium leguminosarum biovar. phaseoli

Rhizobium leguminosarum bv. phaseoli strains P31 and R1, Serratia sp. strain 22b, Pseudomonas sp. strain 24 and Rhizopus sp. strain 68 were examined for their plant growth-promoting potential on lettuce and forage maize. All these phosphate solubilizing microorganisms (PSM) were isolated from Québec soils. The plants were grown in field conditions in three sites having high to low amounts of available P. In site 1 (very fertile soil), strains R1 and 22b tended to increase the dry matter yield of lettuce shoots (p≤0.10). Lettuce inoculated with rhizobia R1 had a 6% higher P concentration (p≤0.10) than the uninoculated control. In site 2 (poorly fertile soil), the dry matter of lettuce shoots was significantly increased (p≤0.05) by inoculation with strain P31 and 24 plus 35 kg ha^-1 P-superphosphate, or with strain 68 plus 70 kg ha^-1 P-superphosphate. In site 3 (moderately fertile soil), the dry matter of maize shoots was significantly increased (p≤0.05) by inoculation with strain 24 plus 17.5 kg ha^-1 P-superphosphate, or with strain P31 plus 35 kg ha^-1 P-superphosphate. Inoculation with PSM did not affect lettuce P uptake in the less fertile soil in site 2. In site 3 with the moderately fertile soil, maize plants inoculated with strain R1 had 8% higher P concentration than the uninoculated control (p≤0.01), and 6% with strains P31 and 68 (p≤0.05). The results clearly demonstrate that rhizobia specifically selected for P solubilization function as plant growth promoting rhizobacteria with the nonlegumes lettuce and maize. The P solubilization effect seems to be the most important mechanism of plant growth promotion in moderately fertile and very fertile soils when P uptake was increased with rhizobia and other PSM.     

Sorption of phosphorus by the iron oxide-impregnated filter paper (Pi soil test) embedded in soils

Incubation experiments were carried out to evaluate the feasibility of extracting phosphorus from soil by embedding iron oxide-impregnanted filter paper strips (P_i strips) in soils having a wide range in pH, texture, and extractable-P contents. Under flooded conditions, the amount of P extracted by the P_i strips increased with the period of submergence and embedding time of the P_i strips. Under unsaturated conditions, the P_i strips were found to extract P from soils over a wide range in moisture conditions; however, keeping the soil at moisture level between saturation and field capacity was found to result in maximal sorption of P by the strips. An embedding time of 16 h was found to be adequate.Phosphorus extracted by embedding P_i strips in soil columns for 16 h at field capacity moisture level correlated significantly with P extracted by shaking the soil with 0.01 M CaCl₂ solution and a P_i strip for 16 h in the laboratory (r=0.94^**). The P extracted by embedding P_i strips correlated best with Bray 1 P in acid soils (r=0.97^**) and with Olsen P in alkaline and calcareous soils (r=0.96^**). The results of the studies demonstrate the feasibility of developing a nondestructive method of monitoring changes in plant-available P in situ under field conditions.     

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

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

Relationships between physiological and morphological attributes related to phosphate uptake in 25 genotypes of Arabidopsis thaliana

The degree of genetic integration among morphological and physiological characters associated with phosphate uptake in impoverished versus fertile sites was studied among 25 inbred homozygous lines of Arabidopsis thaliana (L.) Heynh. The characters recorded were initial uptake rates, C_min (which reflects the ability to deplete phosphate to low concentrations), SRL (specific root length), and root: shoot allocation variables. Highly significant genotypic differentiation was detected for all of the variables (P<0.001). Cmin was correlated with influx per cm or mg of root; the lines better able to pick up Pi at low concentrations generally had slower initial uptake rates per cm or mg of root. This suggests an evolutionary divergence between a character important for nutrient uptake under fertile conditions, and one important for nutrient uptake under impoverished conditions. This relationship however, may not be valid due to the non-proportional relationship between root length or mass and influx. With a measure of influx that took this non-proportionality into account, the relationship became nonsignificant. SRL was negatively correlated (P<0.05) with influx per unit length of root. This suggests that there is an allocation-based trade-off between root length and diameter among Arabidopsis thaliana genotypes. Lower uptake rates per unit length of root for genotypes with thin roots may simply be a consequence of lower root surface area.     

Shading reduces exploitation of soil nitrate and phosphate by Agropyron desertorum and Artemisia tridentata from soils with patchy and uniform nutrient distributions

 Shading may both lessen the demand for soil nutrients and also the energy supply for nutrient acquisition. Since root foraging for nutrients in patchy environments can be energy-costly, especially for an immobile nutrient such as phosphate (P), the effects of shading may be most expected in heterogeneous soils. Plant acquisition of nitrate (N) and phosphate from soils with patchy and uniform nutrient distributions was determined in a field study under open sunlight and with shading for two common perennial Great Basin shrub steppe species, Agropyron desertorum and Artemisia tridentata. Partial shading in a pattern which can occur in shrub steppe vegetation significantly decreased plant N and P acquisition from soils both in the patchy and the uniform nutrient treatments. Artemisia was more affected by the shading than was Agropyron. Exploitation of the rather immobile P ion by both species was reduced to a much greater degree by the shading in the patchy distribution treatment than in the uniform nutrient treatment. As expected, plant acquisition of the more mobile N varied little with nutrient distribution treatment for both species and the depression of N acquisition by shading was the same in both nutrient distributions. The effects of shading appeared to have had its primary influence on different components of root foraging in the two species, especially in the nutrient-rich patches. For Agropyron shading primarily affected root proliferation, as indicated by reduced root density in patches. For Artemisia, shading most influenced root physiological uptake capacity and this was most pronounced in the nutrient-rich patches. While aboveground competition for light may generally reduce nutrient acquisition, the effects appear to be most pronounced if root systems of these steppe species are foraging for nutrients such as P in spatially heterogeneous soils. Received: 29 February 1996 / Accepted: 16 July 1996     

Effects of phosphorus application and mycorrhizal inoculation on root characteristics of subterranean clover and ryegrass in relation to phosphorus uptake

The effects of phosphorus (P) application and mycorrhizal inoculation on the root characteristics of subterranean clover and ryegrass were examined. Phosphorus application increased total root length, root surface area and root volume of both plant species. In contrast, mycorrhizal infection only affected the root characteristics of subterranean clover. Ryegrass took up more P than non-mycorrhizal subterranean clover at all levels of application. However, mycorrhizal infection only increased P uptake by subterranean clover and there was no difference in P uptake between ryegrass and mycorrhizal subterranean clover at low levels of P application. When the P uptake was expressed on the basis of any of the root characteristics, subterranean clover were superior to ryegrass suggesting that the greater uptake of P by ryegrass is not due to a higher efficiency in absorption of P from soil solution, but rather to a large root system.     

Phosphorus Acquisition Strategies within Arbuscular Mycorrhizal Fungal Community of a Single Field Site

Diversity in phosphorus (P) acquisition strategies was assessed among eight isolates of arbuscular mycorrhizal fungi (AMF) belonging to three Glomus species, all obtained from the same field site. Maize (Zea mays L. cv. Corso) was used as a test plant. Compartmented cultivation containers coupled with ³³P radioisotope labeling of soil P were employed to estimate (1) the distance from the roots that AMF were able to acquire soil P from, (2) the rate of soil colonization, (3) the efficiency of uptake of soil P by AMF, (4) benefits provided to maize in terms of P acquisition and growth. Glomus mosseae and G. intraradices took up P 10 cm from roots, whereas G. claroideum only up to 6 cm from the roots. G. mosseae most rapidly colonized the available soil volume and transported significant amounts of P to maize from a distance, but provided no net P uptake benefit to the plants. On the other hand, both G. intraradices and three out of four G. claroideum isolates significantly improved net P uptake by maize. These effects seem to be related to variability between and to a limited extent also within AMF species, in mycelium development, efficiency of hyphal P uptake and effects on plant P acquisition via the root pathway. In spite of absence of maize growth responses to inoculation with any of the AMF isolates, this study indicates remarkable functional diversity in the underground component of the studied field site.