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.     

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.     

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.     

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.     

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.     

Phosphorus Efficiency of Buckwheat (Fagopyrum esculentum)

The ability of buckwheat (Fagopyrum esculentum) roots to acquire phosphorus (P) was characterized by investigating P uptake, morphological features, and chemical changes in the rhizosphere. Over a range of nutrient solution P concentrations (5–500 μmol · L^?1), maximum shoot growth was achieved with a P supply between 5 and 100 μmol · L^?1. Root weight and root length, as well as length and frequency of root hairs, were higher at low P levels. Root surface and the root surface/shoot dry weight ratio reached high values. Though P uptake rates were only moderate (0.15 pmol · cm^?1 root · sec^?1), shoot P concentrations were high (1.8% of dry weight with 100 μM P) predominantly being inorganic (80%). Phosphorus efficiency was characterized by a high specific absorption rate (810 mmol P · kg^?1 root dry wt · d^?1) rather than by an efficient utilization for dry weight production. Root exudates of low-P plants had lower pH values than exudates of high-P plants and increased the solubility of FePO₄ and MnO₂ to a greater extent. Amounts of exuded organic acids and phenolics were low and could not account for the observed solubilization of FePO₄ and MnO₂. Enhanced hydrolysis of glucose-6-phosphate by exudates from low-P plants was due to an increased “soluble” acid phosphatase activity, and root surface phosphatase activity was also slightly enhanced with P deficiency. In the rhizosphere soil of buckwheat, some depletion of organic P forms was observed, and in pot trials with quartz sand, buckwheat utilized glucose- 6-phosphate as a P source at the same rate as inorganic P.     

Einfluss der Kaliumdynamik im wurzelnahen Boden auf die Magnesiumaufnahme von Pflanzen

Influence of potassium dynamics at the soil-root interface on magnesium uptake of plants At the soil-root interface potassium concentration of the soil solution can be depleted by potassium uptake of the plant. The influence of this effect on the magnesium uptake of ryegrass, barley, maize and rape was studied in pot experiments with luvisols from loess. The results have shown that the rate of magnesium uptake was doubled when the potassium concentration at the root surface decreased below 20 μmol K/1. Magnesium uptake is therefore inhibited by K concentrations above this limit. Application of potassium fertilizer increases the potassium concentration of the soil solution. However, via exchange of adsorbed Mg ions from the soil matrix, K application also increases magnesium concentration of the soil solution. As a result of K application magnesium uptake increases in this case if K concentration of the soil solution at the root surface is kept below 20 μmol K/1 by K buffering or K uptake. Magnesium uptake decreases however, if K concentration exceeds 20 μmol/1 soil solution because the inhibitory effect of potassium on Mg uptake is stronger than the favourable influence of Mg concentration.     

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.     

Phosphatverarmung des wurzelnahen Bodens und Phosphataufnahme von Mais und Raps

Phosphate depletion at the soil — root interface and the phosphate uptake of maize and rape Maize and rape plants were grown in flat containers in a ³3P-labelled sandy soil and the distribution of soil phosphate near roots was determined by using densitometric scans of autoradiographs. The concentration of isotopically exchangeable phosphate at the root surface decreased within a few days by 42 per cent with rape and by 50–65 per cent with maize. Initially the width of the depletion zone is very small. Within six days the depletion zone extended to the final distance from the surface of the root cylinders of about 2 mm for maize and 2.6 mm for rape. The soil within the range of the mean length of root hairs (0.7 mm for maize and 1.3 mm for rape) is almost equally depleted. This indicates that root hairs are very important for P-uptake from soil. This is further supported by higher P-uptake rates per cm root length of rape than of maize. The P-concentration of the soil solution was estimated by means of the phosphate desorption curve. Within the root hair cylinder the P-concentration of the soil solution decreased from 0.8 to 0.03 mg P/l. Changes of the P-depletion profile with time were used to calculate P-uptake rates for roots of different age. The results indicate that for the first 3–5 days P-uptake rates remained near maximum, even though the P-concentration of the soil solution at the root surface had strongly decreased within two days. Phosphate uptake rates per cm root length did not decrease unless the whole root hair cylinder had been depleted.     

Potassium efficiency mechanisms of wheat,barley, and sugar beet grown on a K fixing soil under controlled conditions

Plant species differ in their potassium (K) efficiency, but the mechanisms are not clearly documented and understood. Therefore, K efficiency of spring wheat, spring barley, and sugar beet was studied under controlled conditions on a K fixing sandy clay loam. The effect of four K concentrations in soil solution ranging from low (5 and 20 μM K) to high (2.65 and 10 mM K) on plant growth and K uptake was investigated at 3 harvest dates (14, 21, and 31 days after sowing). The following parameters were determined: shoot dry matter (DM), K concentration in shoot dry matter, root length (RL), root length/shoot weight ratio (RSR), shoot growth rate/average root length ratio (GR_s/aRL), K influx, and soil solution K concentrations. Wheat proved to have a higher agronomic K efficiency than barley and sugar beet, indicated by a greater relative yield under K‐deficient conditions. As compared to both cereals, sugar beet was characterized by higher K concentrations in the shoot dry matter, only 30—50 % of the root length, 15—30 % of the RSR and a 3 to 6 times higher GR_s/aRL. This means that the shoot of sugar beet had a 3 to 6 times higher K demand per unit root length. Even at low K concentrations in the soil solution, sugar beet had a 7 to 10 times higher K influx than the cereals, indicating that sugar beet was more effective in removing low available soil K. Wheat and barley were characterized by slow shoot growth, low internal K requirement, i.e. high K utilization efficiency, and high RSR, resulting in a low K demand per unit root length. At low soil K concentrations, both cereals increased K influx with age, an indication of adaptation to K deficiency. The mechanism of this adaptation merits closer investigation. Model calculations were performed to estimate the K concentration difference between the bulk soil and the root surface (ΔC_L) needed to drive the measured K influx. For the two cereals, the calculated ΔC_L was smaller than the K concentration in the soil solution, but for sugar beet, ΔC_L was up to seven times higher. This indicates that sugar beet was able to mobilize K in the rhizosphere, but the mechanisms responsible for this mobilization remain to be studied.     

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.     

Growth and uptake rates of P,K, Ca and Mg in wheat 1

The relation between plant age and nutrient absorption properties of red winter wheat (Triticum aestivum L.) roots were investigated. Understanding the change in ion uptake parameters with increasing plant age is helpful in devising efficient fertilization systems. Such information can be used to determine the nutrient levels needed in the soil to supply nutrients rapidly enough to the root surface to minimize deficiencies. Wheat was grown for periods up to 40 days in solution culture in a controlled climate chamber. Sequential harvest and nutrient influx measurements were made. Shoot growth was exponential with time to 30 days and linear thereafter. Root dry weight increased linearly with time at a slower rate than shoot dry weight. Root length increased linearily with time. With increasing plant age there was a reduction in average P and K uptake rate while average uptake rates for Ca and Mg remained relatively unchanged. With increasing plant age, the maximum influx, I_max. for P and Mg remained constant, but for K and Ca, there was a decrease. For the Michael is constant, Km, no change was observed for P, an increase occurred for K, and a decrease for Ca and Mg, as the wheat plant grew from 5 to 40 days.     

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.     

辣椒丛枝菌根真菌吸磷机制研究

A pot experiment was conducted to investigate the action mechanisms of arbuscular mycorrhizal (AM) fungi in phosphorus (P) uptake of Capsicum annuum L.in a sterilized fossil Oxisol.Three P levels of 0,10 and 200 mg kg-1 soil (P0,P10 and P200,respectively) without and with AM fungal inoculation were applied as Ca(H2PO4)2·H2O.Shoot dry matter yields and shoot P uptake increased significantly (P > 0.05) by the inoculation of AM fungi at P0 and P10.Root length and P concentration in soil solution increased with the inoculation of AM fungi but the root:shoot ratio decreased or remained constant.Around 50% roots of inoculated plants were infected by AM and the external hyphae amounted to 20 m g-1 soil at P10 and P200.The hyphae surface area of the infected root cylinder amounted to 11 and 2 cm-2 cm-2 root at P0 and P10,respectively.The increased P uptake of inoculated plants was mainly because of an up to 5 times higher P influx of the infected root.Model calculations showed that the root alone could not have achieved the measured P influx in both infected and non-infected roots.But the P influx for hyphae calculated by the model was even much higher than the measured one.The P uptake capacity of hyphae introduced in the model was too high.Model calculations further showed that the depletion zone around roots or hyphae was very narrow.In the case of the root only 7% of the soil volume would contribute P to the plant,while in the case of hyphae it would be 100%.The results together with the model calculations showed that the increased P uptake of AM inoculated plants could be explained partly by the increased P concentration in the soil solution and by the increased P absorbing surface area coming from the external hyphae.     

Influence of soil temperature and soil compaction on growth and P uptake of sugar beet

Sugar beet growth is often impaired by cold and compacted soil. The aim of this study was to determine the effect of soil temperature and soil compaction on the growth and function of sugar beet roots. For this purpose a pot experiment with sugar beet (Beta vulgaris) was conducted in a growth chamber in which the soil temperature was kept constant either at 10°C or 20°C and air temperature at 20°C. The soil was uncompacted (1.30 g cm^?3) or compacted to a bulk density of 1.65 g cm^?3. In order to find out whether growth restriction was caused by insufficient P supply of the plant the experiment was run without and with P application (300 mg per kg soil). Root growth was much smaller at 10°C compared to 20°C, whereas root/shoot ratio was not affected by soil temperature. Hence, root and shoot growth was inhibited to the same extent. P content of the plants was not reduced, neither by cold nor by compacted soil, although parameters of acquisition such as root length and morphological root properties were altered. Soil temperature strongly affected P influx, whereas compaction did not. The calculation with a simulation model showed that at 10°C soil temperature the predicted P uptake of the plants agreed with the measured P uptake irrespective of compaction and P application. However, at 20°C the model underestimated the P influx at low soil P availability even if allowance was made for root hairs. It is concluded that under conditions of high shoot P demand and low P availability in soil P has been mobilized by mechanisms not taken into account by the model.     

The acquisition of phosphate by higher plants: Effect of carboxylate release by the roots. A critical review.

Phosphorus is one of the most limiting macronutrients for plant productivity in agriculture worldwide. The main reasons are the limited rock phosphate reserves and the high affinity of phosphate (P) to the soil solid phase, restricting the P availability to the plant roots. Plants can adapt to soils low in available P by changing morphological or/and physiological root features. Morphological changes include the formation of longer root hairs and a higher root : shoot ratio both parameters increasing the root surface which provides the shoot with P. This may be successful if the P availability in soil, i.e., the P concentration of the soil solution is not extremely low (> 1–2 µM P). If the P concentration of the soil solution is lower, the diffusive flux to the root surface will be very low and may not satisfy the P demand of the shoots. Under these conditions plants have developed strategies to increase the rhizosphere soil solution concentration by secreting mobilizing agents. The most effective way of P mobilization is the release of di‐ and tricarboxylic acid anions, especially oxalate and citrate. Citrate can accumulate in the rhizosphere up to concentrations up to 80 µmol g^?1 soil. Cluster root formation is an efficient way of carboxylate accumulation in the cluster root rhizosphere improving P mobilization. Cluster roots strongly improve the acquisition of the mobilized P. Considering a single root, around 80–90% of the mobilized P diffuses away from the root. From the rhizosphere of cluster roots, most of the mobilized P is taken up by the cluster roots. Both, the strong accumulation of carboxylates in and the effective P uptake from the cluster‐root rhizosphere are the basis of the unique ability of P acquisition by cluster root‐forming plants. Plants that do not form cluster roots, e.g., red clover, can also accumulate carboxylates in the rhizosphere. Red clover accumulates high quantities of citrate in the rhizosphere soil. Model calculations show that the release of citrate by red clover roots and its accumulation in the rhizosphere strongly improve P acquisition by this plant species in various soils. Similar results are obtained with alfalfa. In sugar beet, oxalate release can strongly contribute to P acquisition. In summary, P acquisition can be strongly improved by the release of carboxylates and should be taken as a challenge for basic and applied research.     

Phosphorus efficiency of ornamental plants in peat substrates

A pot experiment was conducted to investigate factors contributing to phosphorous (P) efficiency of ornamental plants. Marigold (Tagetes patula) and poinsettia (Euphorbia pulcherima) were cultivated in a peat substrate (black peat 80% + mineral component 20% on a volume basis), treated with P rates of 0, 10, 35, 100, and 170 mg (L substrate)^–1. During the cultivation period, plants were fertigated with a complete nutrient solution (including 18 mg P L^–1) every 2 d. Both poinsettia and marigold attained their optimum yield at the rate of 35 mg P (L substrate)^–1 and the critical level of P in shoot dry matter of both crops was 5–6 mg g^–1. After planting, plant‐available P increased at lower P rates to a higher level for poinsettia than for marigold, but no significant change was observed at higher P rates. Balance sheet calculations indicated that at lower P rates more P was fertigated than was taken up by the plants. Root‐length density, root‐to‐shoot ratio, and root‐hair length of marigold were doubled compared to that of poinsettia. Root‐length density increased with crop growth, and 10 d after planting the mean half distance between roots exceeded the P‐depletion zone around roots by a factor of 3 and 1.5 for poinsettia and marigold, respectively. Thus, at this early stage poinsettia exploited only 10% of the substrate volume whereas marigold utilized 43%. Later in the cultivation period, the depletion zones around roots overlapped for both crops. Taking into account P uptake via root hairs, the simulation revealed that this was more important for marigold compared to poinsettia especially at low P‐supply levels. However, increase of P uptake due to root hairs was only 10%–20% at optimum P supply. For the two lower P levels, the P‐depletion profile around roots calculated for 10 d after planting showed that after 2 d of depletion the concentration at the root surface was below the assumed K_m value (5 μM) and the concentration gradient was insufficient to fit the demand. A higher content of plant‐available P in the substrate was observed for poinsettia compared to marigold in the treatment with P application adequate for optimum growth, because more fertigated P was accumulated during early stages of cultivation due to lower root‐length density of poinsettia. The observed difference of root morphological parameters did not contribute significantly to P‐uptake efficiency, since P mobility in the peat substrate was high.     

Growth and nutrient uptake by cotton roots under field conditions

Abstract

The relationship between nutrient uptake and root growth of cotton (Gossypium hirsutum L.) was studied under field conditions. This basic information could be beneficial when making best management decisions concerning the time of application and placement of fertilizer. A field study was conducted in North Alabama on a fertile Dewey silt loam (clayey, kaolinitic, thermic Typic Paleudult). Aboveground whole plants were harvested at approximately 10‐day intervals beginning at 211 cumulative heat units (CHU) after planting (37 days after planting: 4‐true leaves). Root length of harvested plants was also measured by depth and distance from the plant. Maximum root length was obtained at 1174 CHU (117 days after planting), while dry matter continued to increase until a maximum was obtained at 1317 CHU (128 days after planting). Maximum root length density of 1.60 cm cm³ was obtained in the surface 0–15 cm layer in the in‐row position at 912 CHU (99 days after planting). After first bloom approximately 70% of the cotton root system was in the surface 30 cm of soil. Average daily influx of S per m of root length increased with plant age until 1317 CHU (near cut‐out), after which influx declined. Nitrogen (N), calcium (Ca), and iron (Fe) influx peaked very early in the season (291–469 CHU) followed by a general decrease with plant age. Maximum daily influx of potassium (K), phosphorus (P), magnesium (Mg), copper (Cu), manganese (Mn), and zinc (Zn) per meter of root occurred at approximately peak‐bloom (764–912 CHU, 87–99 days after planting) and decreased with plant age. Copper, Fe, Mn, and Zn influx rates were ～ 1000 times lower as compared to the other nutrients.