Wurzelstudien und Phosphat-Aufnahme von Weidelgras und Rotklee unter Feldbedingungen

Root studies and phosphorus uptake of rye-grass and red clover under field conditions Root parameters (fresh weight, density, surface, length, cation exchange capacity) and phosphate uptake were studied with rye grass and red clover, grown in the field on a brown podsolic soil. In all root parameters, ry grass was superior to red clover. Also, phosphate uptake of rye grass was higher than that of red clover. The greatest difference between both species was found in root length, that of rye grass being about five times longer than that of red clover. Rye grass had longer root hairs than red clover; whereas root diameter of clover was about twice as thick the average rye grass. Significant correlations were observed between root parameters and phosphate uptake in the plants studied. The highest correlation coefficients were obtained for the relationship P-uptake versus root length (clover 0.91***, grass 0.87***) and P-uptake versus root fresh weight (clover 0.92***, grass 0.88***). The phosphate uptake per unit root parameters was significantly higher in red clover, compared with rye grass, for the parameters root fresh weight, cation exchange capacity and root length. Because of this high P-uptake rate for clover it is assumed that clover also requires a higher P-concentration in soil solution as compared with grass. Thus grass may still grow with low P concentrations in the soil solution without P deficiency at which clover cannot grow. It is for this reason that in mixed swards clover is depressed by grass, if the available P in the soil is low.     

Wirkung der Kaliumdüngung auf die Kaliumverfügbarkeit in der Rhizosphäre von Raps

Influence of potassium supply on the availability of potassium in the rhizosphere of rape (Brassica napus) Potassium depletion of the soil in the proximity of roots was studied in order to obtain quantitative information on the availability of potassium. For this purpose rape seedlings were grown in pots which separate roots from soil by a fine meshed screen; root hairs penetrated the screen. The soil adjacent to the screen was sliced by microtome into layers about 0.1 mm thick which were separately analysed for k. Plant roots strongly depleted the soil in their proximity; further distant ranges remained unchanged. A loess derived loam, brought to different levels of exchangeable K by precropping or K application, was equally depleted to 150 μmoles/100g soil at the root surface. Therefore, the quantity of K released from this source increased with initial K level. In addition, the distance of the depletion zone extended with K level from 4,6 to 6.3 mm from the surface of the root cylinder. Hence, the volume of soil contributing K to the root increased from 0.7 to 1.2 cm³ per cm root length. The combination of these two parameters, i.e. the quantity of exchangeable soil K released per unit of root length increased by a factor of 20 whereas exchangeable soil K was raised by a factor of 4.5 only. K uptake of the plants after 4 days was in agreement with the depletion of exchangeable soil K in the high K treatment only. The other treatments had obviously taken up considerable percentages of nonexchangeable K. This fraction was released from the soil ranging less than 1.5 mm from the root. The distance of the K depletion zone was also extended by application of NaCl and MgCl². Because of cation exchange, K concentration of the soil solution was increased, K buffer power decreased and, therefore, K diffusion was enhanced. It is concluded that plants in the field do not uniformly deplete the total rooted soil volume. Whereas roots in their proximity strongly deplete the soil including parts of nonexchangeable K they do not even use exchangeable K in a slightly greater distance. The quantity of K available per unit of root length is, therefore, determined by both - the degree of soil K depletion at the root surface and -the distance of the depletion zone, i.e. the volume of soil that contributes K to the root. Either factor was markedly affected by the level of soil K and thus by K application.     

Kaliumdynamik im wurzelnahen Boden in Beziehung zur Kaliumaufnahme von Maispflanzen

Potassium dynamics at the soil-root interface in relation to the uptake of potassium by maize plants Young maize plants were grown in flat containers on a sandy and a silt loam soil after addition of ⁴³K as tracer. Changes of the K concentration in soil in the vicinity of the roots were determined by scanning the film density of autoradiographs. A distinct zone of K depletion in the soil adjacent to the root surface was observed, similar to those found earlier with phosphate and rubidium. The highest degree of depletion occured within a distance of 0.7 mm from the surface of the root cylinder which corresponds to the average length of root hairs of the cultivar used. The quantity of K released within 2.5 days per unit of this part of the soil exceeded the exchangeable K by a factor of two. In a radial direction the zone of maximum depletion was followed by a depletion profile which extended over 5 mm in the sandy and over 3 mm in the silt loam soil. The K concentration of the soil solution decreased to 2–3 μmoles K/l at the root surface. In order to determine the effect of depleting the K concentration by plant roots on the release of soil K, desorption studies were carried out in parallel. For this purpose the soil was successively extracted by solutions with cation concentrations corresponding to the soil solution, except for K. With this procedure a massive release of K from the soil was observed after the equilibrium concentration decreased to 2–3 μmoles K/l. It is concluded that

• – in one growing season only part of the soil volume of the rooted layer contributes potassium to the plant and, on the other hand
• – substantial part of the potassium absorbed by plants is derived from nonexchangeable soil K, even in short periods of time.

     

Bedeutung von Kaliumaufnahmerate,Wurzelwachstum und Wurzelhaaren für das Kaliumaneignungsvermögen verschiedener Pflanzenarten

Effect of K uptake rate, root growth and root hairs on potassium uptake efficiency of several plant species Pot experiments with maize, rape, tomato, rye-grass and onion plants were carried out to evaluate the influence of – rate of K uptake per cm of root, – cm root per mg shoot dry weight and – mean root age (as a measure of the time roots absorb potassium) on potassium uptake efficiency of these plants. Percent K in shoot dry matter was used to indicate K uptake efficiency. No close correlation was observed between one of these factors to K concentration in shoot dry matter. The product of K uptake rate and root-shoot ratio was closely related to the K concentration of shoots. However, regression lines for maize, rape and onion were different. One single regression line was found when K concentration in shoot was related to the product of K uptake rate, root-shoot ratio and mean root age. It is therefore concluded that K uptake of plants depends on all three of these factors. In different species the proportion of these factors were markedly different. The plant factors in turn were affected by the K nutritional status of the plants. K uptake rate increased whereas root-shoot ratio and mean root age decreased with increasing K supply of the soil. K uptake rate per cm root was strongly affected by root hairs. The radial distance of the K (Rb) depletion zone of the soil adjacent to the root surface also increased with the length of the root hairs. It is therefore concluded that root hairs substantially affect the spatial access of potassium in soil by the plant.     

INTERPRETATION OF PHOSPHORUS UPTAKE BY BARLEY FROM ACID SOILS USING A DIFFUSION MODEL

Phosphorus uptake by bailey (per cent of dry tissue) grown on four acid soils, each treated with four amounts of lime and P, correlated equally well with EPC (equilibrium P concentration = concentration of P supported in solution at field capacity) and with diffusion supply of P to root cylinder surfaces calculated from the soil indices including EPC, a soil capacity factor maintaining that concentration, and a diffusion coefficient. Calculated rates of P supplied by diffusion to primary root cylinder surfaces were too low to account for P found in plants. If the remainder of the observed P uptake was assigned to root hairs, the ratios of calculated root hair surface area to estimated root cylinder surface areas necessary to account for the difference were in reasonable agreement with ratios for barley reported by other workers.     

Rubidium-Verarmung des wurzelnahen Bodens durch Maispflanzen

Rubidium depletion of the soil-root interface by maize plants Maize plants were grown in flat containers with radioactive labelled rubidium. Changes of the Rb concentration in soil in the vicinity of the roots were determined by means of the film density of autoradiographs. Results were as follows: The Rb concentration of the soil at the root surface decreased markedly within one day; only small changes occured after this period. Initially, the width of the depletion zone was very small. It extended in the following days in a radial direction. Therefore, after the initial phase the Rb supply of the plants depended on transport from more remote parts of the soil. Soil texture and Rb level strongly influenced both degree and distance of Rb depletion. Thus, the Rb concentration at the root surface decreased by 80% of the initial value in a sandy soil (4% clay) and by only 30% in a silt loam soil (loess, 21% clay). The depletion zone extended to a distance of 2 mm in the silt loam soil from the surface of the root cylinder and to 5 mm in the sandy soil. Hence, in the silt loam about 20% and in the sandy soil almost 100% of the total soil volume contributed Rb to the plant, assuming a root density of 1 cm per cm³ of soil. Increased levels of Rb enhanced Rb availability by increasing both the degree of soil depletion near the root surface and the size of the depletion zone. The quantity of Rb available per cm of root varied between 0.05 μmol in the silt loam with low Rb application and 2.7 μmol in the sandy soil with high Rb application. The amount of Rb depleted from the soil, expressed as per cent of the Rb exchangeable by ammoniumacetate ranged from 3 to 7% in the silt loam and from 20 to 30% in the sandy soil, calculated on the basis of 1 cm root per cm³ of soil. The Rb concentration of the soil solution near the root surface was reduced to 2 μmolar.     

Availability of phosphate and potassium as the result of interactions between root and soil in the rhizosphere

A number of findings are summarized in order to show the significance of individual plant properties and soil factors on the availability of phosphate and potassium to plants growing in soil. The flux of a nutrient into a given plant root depends directly on the concentration of the nutrient in the adjacent solution. In nutrient solution, P and K influx follows Michaelis-Menten kinetics. Almost maximum rates of influx have been observed in the range of soil solution concentrations usually found in German arable soils. Roots exhaust P and K from solutions to about 0.2 μmol P and 1 μmol K 1^?1 if not replenished. At the root surface P and K concentrations in soil decrease rapidly within one day; small changes occur after this period. Initially, the extent of the depletion zone is very small but it extends radially with time. After the initial phase therefore, P and K supply to the plant depends on transport from more remote parts of the soil and also on release from undissolved sources. The degree of depletion and the extent of the depletion zone are related to the diffusion coefficient; they decrease with increasing clay content of soil. Root hairs penetrate the soil and extend the volume of soil supplying nutrients to a unit of root. P and K influx therefore increase with the length of root hairs. Proton release of roots mobilize P and K in soil. This is clearly detected by the HCl-soluble P and K fractions within 2 mm of the root surface. The activity of acid and alkaline phosphatases strongly increase in the soil in the vicinity of the root surface of several plant species. It is supposed that organic P compounds can therefore be utilized by plants. P and K influx per unit of root length and root length per unit of shoot weight differed widely between species. The product of these two parameters however was closely related to the P and K concentration of the shoots. Calculations from a mathematical model were in good agreement with measured K depletion profiles and K uptake by plants. It is therefore concluded that the main factors influencing the P and K availability of plants growing in soil have been accounted for in the mathematical model and that the parameters have been accurately measured.     

Erfassung der Mineralstoffverteilung in Wurzelnähe durch getrennte Analyse von Rhizo- und Restboden

Estimation of the nutrient distribution around plant roots by separate analysis of rhizo and bulk soil A procedure is described to separate rhizo soil from bulk soil. For this purpose the living plant roots along with the strongly adhering soil are taken out of the surrounding soil. After a short period of drying, roots are separated from rhizo soil by gently sifting. Chemical analysis of the samples has shown that the nutrient content of rhizo soil usually differs markedly from bulk soil. Phosphate and potassium content of the rhizo soil of wheat and maize is lower than in bulk soil. Percentagewise, these differences are higher by using water as an extractant when compared to the slightly acid lactate or formate solutions. The water soluble calcium content of several wheat fields was in most cases higher in the rhizo than in the bulk soil. In sandy soil the depletion zone of phosphate does not exceed the length of the root hairs markedly whereas in the case of potassium it does. This is concluded from data showing that the potassium content, but not the phosphate content, was decreased in the bulk soil if compared to the uncropped soil. In heavier loess soils the nutrient concentrations of rhizo and bulk soil were not always found to be markedly different. It is assumed that the depletion zones are smaller than in sandy soils. In addition, because of aggregation, rhizo soil cannot be separated from bulk soil as neatly as in sandy soils. Therefore, in heavier soils the method does not yield satisfactory results.     

The maize root system in situ: Evaluation of structure and capability of utilization of phytate and inorganic soil phosphates

• 1 The dependence of the morphology of the maize (Zea mays L.) seminal root system on physical, chemical and biotic parameters was investigated with pot cultures in quartz sand and in a natural loamy sand soil. Low O₂-supply to the soil resulted in a substantially smaller root biomass despite a relative increase in total root length. Reduced N-supply also stimulated root length growth, but also enhanced the formation of laterals. The presence of soil microorganisms, in comparison to sterile cultures, resulted in a reduced length of the main roots, and the production of slender laterals with a decreased root hair density. Generally, the structural variability of laterals in response to different growth conditions was much more pronounced than that of the main roots.
• 2 A major part of the work reported here was dedicated to a detailed study of phosphate (P) acquisition by the maize root system under field conditions. Radioactive labelling of the roots and radioautography of soil cores revealed the in situ distribution pattern of the maize root system. Controlled labelling of the soil with radioactive phosphate allowed the documentation of the development and replenishment of the phosphate depletion zone around roots. Finally, the longevity and phosphate uptake activity of the different parts and tissues of the primary root system of maize was examined by electron microscopy and tracer studies including pulse chase experiments. From these studies the phosphate-acquiring strategy of the maize root system appears as follows: The capability of P uptake decreases in the order: root hairs, 1^st order laterals, 2^nd order laterals, main root. The life-spans of the components of the maize root system increase by the sequence: root hairs, laterals, main root. Inorganic P uptake, therefore, mainly occurs during the first weeks of root development. Dying back of the root occurs in an ordered manner resulting in a relocation of stored P predominantly into the main root cortex. Furthermore, it could be shown that competition for P between roots of the same or of adjacent maize and/or lupin plants virtually does not occur in situ.
• 3 The utilization of phytate-P was studied with ¹⁴C/³²P-labelled Camyo-inositol-hexaphosphate supplied to maize plants grown in sterile quartz sand or in hydroponic cultures. The ratio of P- and C-uptake as well as the incidence of phytate hydrolysis products in the rooting medium indicated the capability of maize roots to acquire P from phytate by enzymatic hydrolysis. This was confirmed by enzyme studies of the root tissues. A specific hydrolyzing enzyme (phytase; molecular weight 51 kD) could be detected in the cell wall of the root, especially in the root tip, which initiates phytate dephosphorylation. Further breakdown is presumably accomplished by monophosphoric phosphohydrolases.

     

Estimation of phosphorus uptake capacity by different zones of the primary root of soil-grown maize (Zea mays L.)

A method is described which allows determination of the nutrient uptake capacity by different zones of individual roots of soil-grown plants. Examples are given for phosphorus uptake by different zones of the primary root of maize. Agar strips with labelled phosphorus (³²P) are placed on the soil-root interface of root zones of different age. After 24 h autoradiographs are made of the soil-root interface and phosphorus uptake rates determined by plant analysis. Along the primary root the relative phosphorus uptake capacity declines per unit root length from 100% in 1 day old root zones to about 25–30% in 26 day old root zones. The change in the extension of the phosphorus (³²P) depletion zone along the root axis indicates that the uptake capacity for phosphorus of root hairs was maximal upto 4 days and declined thereafter. The decline in capacity for phosphorus uptake from apical to basal root zones probably reflects the decline in root hair viability. The relative high uptake capacity in basal root zones is therefore at least partially due to the uptake capacity of the cortical cells.     

Availability in Soil and Acquisition by Plants as the Basis for Phosphorus and Potassium supply to Plants

This report summarizes research aimed at describing the processes and quantifying the factors affecting transfer of P and K from soil into plants. Soil properties related to availability and plant properties reflecting nutrient acquisition were determined. Their interactions in the rhizosphere and their importance for nutrient supply of plants were studied by a combination of measurements and calculations using a simulation model. Phosphorus and potassium uptake by roots decreased P and K concentration at the root surface and caused characteristic depletion profiles in the adjacent soil. The shape of the profiles depended on the effective diffusion coefficient, the concentration of the nutrient in soil, morphological properties of the roots and on influx into roots. The degree of depletion at the root surface indicated the proportion of the nutrient potentially available in the soil. The shape of the depletion profiles reflected the amount of the nutrient taken up by a root section. The parameters found to describe nutrient acquisition are (i) influx per unit root length, (ii) root length per unit shoot weight (root/shoot ratio), and (iii) the period of time a root section absorbs nutrients. Plant species differed considerably in these properties. In order to integrate the processes involved and to evaluate the importance of individual factors, the Claassen-Barber model was used. Depletion profiles and nutrient uptake calculated with this model were in good agreement with measured values in a number of cases. However, at low P supply, plants absorbed substantially more P than the model predicted. This indicates that influx in this case is supported by mechanisms not properly taken into account yet. Influx per unit root length depends on morphological properties of and nutrient mobilization by roots. Root hairs increase root surface area per unit root length. In addition, because of their small diameter and geometric arrangement in soil, root hairs are specially apt to gain from diffusion when concentration gradients are small. This applies even more to VA-mycorrhizae. Their hyphae are longer and thinner than root hairs and can thus deplete larger volumes of soil per unit root length. Root-induced changes of soil pH increased the size of P depletion profiles, indicating that roots can mobilize soil P by this mechanism. Both acid and alkaline phosphatase enzyme activities were found to be markedly increased at the soil-root interface suggesting that soil organic P may contribute to the P supply of plants.     

Einfluß der P-Aufnahme und P-Remobilisierung in der Kornfüllungsperiode auf den Ertrag von Sommerweizen

Influence of P-uptake and P-remobilization during the grain-filling period on the yield of spring wheat Experiments were performed with spring wheat in hydroponics using labelling with ³³P and ³²P to investigate the influence of high (0,5 mM) or low (0,05 mM) P-concentration in the solution on the distribution and remobilization of the nutrient as well as on crop yield. With low P-supply the yield was reduced, not only due to a considerable decrease of tillers but also in consequence of lowering the thousand grain weight and the number of grains even in apparently still fully developed ears, as well as significantly depressing the P-concentration in the grain. Flowering and maturity commenced much earlier than in the case of plants supplied with a relatively high level of phosphorus. When the P-concentration in the medium was only dropped from a high to low level after the end of flowering, the yield of the plants, which had a comparable number of tillers to the control, was slightly but significantly reduced, too. In spite of an enormous enhancement of the retranslocation of P deposited in the plant before flowering the P-concentration of the grain was also deminished. The experiments have shown that the intensified P-retranslocation during the grain-filling period and a still continuing P-uptake and translocation into the leaves and particularly the grain proceeded simultaneously under the experimental conditions. Besides, the effective P-retranslocation based on the labelling was substantially greater than indicated by the net P-retranslocation. The possible function of phosphate during the storage phase of cereals is discussed.     

Root trait plasticity and plant nutrient acquisition in phosphorus limited soil

To overcome soil nutrient limitation, many plants have developed complex nutrient acquisition strategies including altering root morphology, root hair formation or colonization by arbuscular mycorrhizal fungi (AMF). The interactions of these strategies and their plasticity are, however, affected by soil nutrient status throughout plant growth. Such plasticity is decisive for plant phosphorus (P) acquisition in P‐limited soils. We investigated the P acquisition strategies and their plasticity of two maize genotypes characterized by the presence or absence of root hairs. We hypothesized that in the absence of root hairs plant growth is facilitated by traits with complementary functions, e.g., by higher root mycorrhizal colonization. This dependence on complementary traits will decrease in P fertilized soils. At early growth stages, root hairs are of little benefit for nutrient uptake. Regardless of the presence or absence of root hairs, plants produced average root biomass of 0.14 g per plant and exhibited 23% root mycorrhizal colonization. At later growth stages of maize, contrasting mechanisms with functional complementarity explained similar plant biomass production under P limitation: the presence of root hairs versus higher root mycorrhizal colonization (67%) favored by increased fine root diameter in absence of root hairs. P fertilization decreased the dependence of plant on specific root traits for nutrient acquisition. Through root trait plasticity, plants can minimize trade‐offs for developing and maintaining functional traits, while increasing the benefit in terms of nutrient acquisition and plant growth. The present study highlights the plasticity of functional root traits for efficient nutrient acquisition strategies in agricultural systems with low nutrient availability.     

Low-P solution culture can be used for screening root growth vigor in soil for high nutrient uptake of spring wheat varieties

Purpose: Root and root hairs of plants have been intensively studied in solution culture; however, correlation of such measurements in solution culture with development in soil is poorly understood. Therefore, the aim of this study is to study whether root and root hairs grown in solution culture can predict their behavior in soil and their correlation with macro- and micronutrients uptake of wheat genotypes.

Materials and methods: The growth of roots and root hairs as well as uptake of macro- and micronutrients of six spring wheat varieties was compared in solution culture under P stress and P abundance and in a low fertility soil.

Results and conclusions: Root length and surface area under P stress were significantly positively correlated with that in the low fertility soil, while no such correlation was apparent for root hair length and density. In absolute terms, the root length, surface area, root hair length and density of spring wheat varieties were substantially higher in soil than in solution culture, while the concentration and uptake of macro- and micronutrients in soil differed from solution culture in a complex way. The early uptake of macro- and micronutrients was intimately associated with root length and surface area as well as root hair length and density in soil but not in solution culture. Therefore, root length rather than root hair traits in low-P solution may be used to screen early root growth vigor in soil and thereby high nutrient uptake of wheat in low fertility soil.     

Improved phosphorus efficiency of three new wheat genotypes from CIMMYT in comparison with an older Mexican variety

In a field trial in Northwest Mexico, the phosphorus efficiency of three advanced bread wheat lines (Triticum aestivum L.) from CIMMYT were compared with an older Mexican variety Curinda, under irrigation, on an alkaline clay soil (3.7 mg Olsen-P kg^—1 soil) without (P-0) and with P-fertilisation (P-35; 35 kg P ha^—1). Dry matter, P-content, P-uptake of above ground biomass and root growth (root length densities in different soil depths) were measured at different growth stages, and the net P-uptake rates per unit root length calculated. All four genotypes responded positively to P-fertilisation. The three new genotypes showed significantly higher grain yields compared with the old variety Curinda, on the average, 54% and 42% higher at P-0 and P-35, respectively. The higher grain yield was mainly due to a larger number of kernels per ear, higher thousand kernel weight as well as a higher harvest index. The old variety Curinda had the same (P-0) or greater (P-35) number of spikes m^—2 than the new genotypes. In conclusion of this experiment, the three new genotypes could be classified as more P-efficient. The P-uptake at harvest averaged 35% and 24% more than the old variety Curinda at the P-0 and P-35 level, respectively. The improved P-efficiency was mainly due to a more efficient P-uptake. However, there were only small differences in P-utilisation efficiency (kg grain per kg P in shoots) between old and new varieties (8—11%). The differences in the root systems were more decisive in the P-0 treatment than with P-fertilisation. At low P, the improved P-uptake per ha of the advanced lines was due to a higher root length density especially after flowering, while at high P, a higher P-influx rate per unit root length played a more important role than the root length density. The superiority of the new genotypes at both P levels is obviously due to the good adaptation of their root system (root length density, uptake rate per unit root) to variable P availability in soil.     

Comparison of root and root hair growth in solution and soil culture

Aerated solution culture is frequently used for studying plant growth. Few comparisons have been made of root growth in solution with that found in soil. The objective of this study was to compare root growth and root hair development in these two mediums. Corn (Zea mays L.) grown in aerated solution at two temperatures (18 and 25°C) and three P concentrations (2, 10, and 500 μmol L^‐1) was compared with that in three soils, Raub (Aquic Argiudoll) and two Chalmers (Typic Haplaquoll) silt loams, in a controlled climate chamber over 21d. Corn plant weight and root growth were similar in solution culture and Raub soil when grown at an air and soil temperature of 18°C. At 25°C both yield and root growth were greater in Raub soil, even though P uptake by corn was 7‐fold greater in solution culture. The same difference was found when corn grown at 25°C in solution culture at 3 different P concentrations was compared with that grown in Chalmers soil at two P levels. Percentage of total root length with root hairs, root length and density and consequently root surface area, were all greater in the Chalmers soil than in solution culture. An increase in soil P, resulted in a decrease in root hair growth. No such relationship was found in solution culture. Although the recovery and measurement of plant roots and root hairs is more convenient in solution culture, results from this study indicate that the usefulness of solution culture for determining those factors which control root growth and root hair development in soil is limited.     

Root hairs and the acquisition of plant nutrients from soil

The literature on the role of root hairs when plants acquire mineral nutrients from soil is reviewed. After a short outline of the root properties affecting the acquisition of nutrients, the roles of root hairs are discussed in four sections, entitled: morphological properties of root hairs, mode of action of root hairs, factors affecting the formation of root hairs, and relationship between root hair formation and plant nutrient uptake. The formation of root hairs depends on both genetic and environmental factors, particularly the supply of phosphate and nitrate. It is concluded that root hairs may substantially contribute to the acquisition of nutrients, mainly those of low mobility in soil and high demand in plants. The percentage of a nutrient acquired by root hairs varies widely, from almost zero to approximately 80 % of the total uptake of the nutrient. The contribution of root hairs depends on plant species and the genetic variability of root hair formation on the one hand, and the kind of nutrient and its availability in soil on the other. According to the published reports, essentially only phosphorus and potassium were considered.     

根-土界面区域磷酸盐的分布和移动: Ⅲ. 动力学

The depletion rate of phosphate in the soil-root interface zone increased along with growth and phosphate uptske of wheat or maize,which indicated that the phosphate distribution in soil near the root surface agreed well with the phosphate movement in rhizosphere and phosphate uptake by plant,The relative accumulation zone of phosphate within 0.5mm apart from the root surface developed at the 15th day or so after cultivating wheat or maize since the root phosphate secretion increased gradually in this stage.The phosphate distribution in the soil-root interface zone against the growing time(t)and the distance from the root plane(x) could be described by the non-linear regression equation with the third powers of x and t.     

Der Einfluß von Bodenart,Durchlüftung des Bodens,N-Ernährung und Rhizosphärenflora auf die Morphologie des seminalen Wurzelsystems von Mais

Influence of soil type, soil aeration, nitrogen supply and rhizosphere flora on the morphology of the seminal root system of maize The influence of the soil type (quartz sand – humous loamy sandy soil), soil aeration, nitrogen supply and rhizosphere flora on the morphology of the seminal root system of maize plants grown in pot culture was investigated. The morphological parameters of number, length, diameter and root hair formation (both length and density) of the main and the lateral roots were determined in addition to the total root length and number and the lateral root density. 1. The biomass production of the shoot and root system was nearly identical in both soils. The total root length growth, however, was enhanced in the sandy soil due to the stimulated formation of first order lateral roots. This increase was correlated with a decrease in the mean diameter and root hair length of the main and lateral roots. 2. A decreased O₂-supply to the soil resulted in a drastic reduction of root biomass, which was correlated, however, with a (relative) increase in total root length (due to the stimulation of the length growth of the first order lateral roots). The root hair length, on the other hand, was reduced under O₂-deficiency. 3. Reduced N-supply resulted in a decrease of the shoot/root-ratio with both substrates which could be ascribed to the enhanced formation and length of the first order lateral roots. 4. The presence of soil microorganisms in quartz sand culture resulted in a reduction of shoot biomass. In comparison with the sterile control culture the total length of the main roots was retarded, the main and lateral roots were more slender and root hair formation was reduced. 5. The experimental results show that the lateral root system demonstrates a significantly greater plasticity than does the main root system.     

Growth and phosphorus uptake of maize cultivated alone,in mixed culture with other crops or after incorporation of their residues

Intercropping or rotating of P‐efficient crop species which mobilize sparingly soluble P by their root exudates can have beneficial effects on growth and P uptake of P‐inefficient species. We aimed at studying the effect of intercropping or incorporating of crop residues of P‐efficient crops on the components of maize P‐uptake, i.e. the root‐system size and P influx (P‐uptake rate per unit root length). This was studied in 3 pot experiments in a low‐P sandy soil. In the first experiment, maize was intercropped with white lupine, sugar beet or oilseed rape, and with groundnut in the second experiment. In the third experiment, maize was grown after incorporating the crop residues of white lupine, sugar beet or oilseed rape. Maize growth and yield was strongly inhibited when intercropped with white lupine, sugar beet or oilseed rape, probably because of competition for nutrients. But with groundnut as the accompanying species, maize yield was increased by a factor of 3, mainly because of an enhanced P influx. Crop residues of oilseed rape and sugar beet increased the yield of maize by factors 2 and 1.6, respectively, because of a 3 and 2 times higher P uptake as compared to maize grown after maize without incorporation of crop residue. The reason for the higher maize P‐uptake after oilseed rape was an 11 times higher P influx as compared to maize without crop residues, and after sugar beet residues because of an enhanced root growth and a 4 times higher P influx. Lupine residues did not improve maize growth, mainly because of a low P influx, which was even less than that of maize grown without crop residues. The soil solution P concentration and calcium acetate lactate‐extractable P (CAL P) measured in this study did not reflect the P availability as indicated by the plants (P uptake, P influx). This indicates that other mechanisms such as P mobilization in the rhizosphere by root exudates or cell‐wall components were responsible for the increased P availability. These mechanisms need further investigation.