Differences in Root Exudation Among Phosphorus‐Starved Genotypes of Maize and Green Gram and Its Relationship with Phosphorus Uptake

Abstract

Availability of phosphorus (P) in soil and its acquisition by plants is affected by the release of high and low molecular weight root exudates. A study was carried out to ascertain the qualitative and quantitative differences in root exudation among the genotypes of maize (Zea mays L.) and green gram (Vigna radiata L.) under P‐stress. Results showed that both inter‐ and intra‐species differences do exist among maize and green gram in terms of root exudation, P uptake, and shoot and root P content. In general, green gram, a legume crop, had greater root exudation compared to maize. However, the amino acid content of the total root exudates in maize was two‐fold as compared to green gram. The maize and green gram genotypes possessed genetic variability in root exudation. Irrespective of the species or genotypes, a positive relationship was found among P uptake rates, total root exudation, and shoot and root ³²P content. The amount of sugars and amino acid present in the root exudates of P‐starved seedlings also add to the variation in P uptake efficiency of genotypes.     

Phosphorus Influx and Root‐Shoot Relations as Indicators of Phosphorus Efficiency of Different Crops

Abstract

The large variation in phosphorus acquisition efficiency of different crops provides opportunities for screening crop species that perform well on low phosphorus (P) soil. To explain the differences in P efficiency of winter maize (Zea mays L.), wheat (Triticum aestivum L.), and chickpea (Cicer arietinum L.), a green house pot experiment was conducted by using P‐deficient Typic ustochrept loamy sand soil (0.5 M NaHCO₃‐extractable P 4.9 mg kg^?1, pH 7.5, and organic carbon 2.7 g kg^?1) treated with 0, 30, and 60 mg P kg^?1 soil. Under P deficiency conditions, winter maize produced 76% of its maximum shoot dry weight (SDW) with 0.2% P in shoot, whereas chickpea and wheat produced about 30% of their maximum SDW with more than 0.25% P in shoot. Root length (RL) of winter maize, wheat, and chickpea were 83, 48, and 19% of their maximum RL, respectively. Considering relative shoot yield as a measure of efficiency, winter maize was more P efficient than wheat and chickpea. Winter maize had lower RL/SDW ratio than that of wheat, but it was more P efficient because it could maintain 2.2 times higher P influx even under P deficiency conditions. In addition, winter maize had low internal P requirement and 3.3 times higher shoot demand (i.e., higher amount of shoot produced per cm of root per second). Even though chickpea had 1.2 times higher P influx than winter maize, it was less P efficient because of few roots (i.e., less RL per unit SDW). Nutrient uptake model (NST 3.0) calculations satisfactorily predicted P influxes by all the three crops under sufficient P supply conditions (C_Li 48 µM), and the calculated values of P influx were 81–99% of the measured values. However, in no‐P treatment (C_Li 3.9 µM), under prediction of measured P influx indicated the importance of root exudates and/or mycorrhizae that increase P solubility in the rhizosphere. Sensitivity analysis showed that in low P soils, the initial soil solution P concentration (C_Li) was the most sensitive factor controlling P influx in all the three crops.     

Impact of fresh root material and mature crop residues of oilseed rape (Brassica napus) on microbial communities associated with subsequent oilseed rape

Glasshouse bioassays were conducted to assess the impact of different inputs of oilseed rape plant material on soil and rhizosphere microbial diversity associated with subsequently grown oilseed rape (Brassica napus) plants. The first bioassay focussed on the effect of oilseed rape rhizodeposits and fresh detached root material on microbial communities, in a rapid-cycling experiment in which oilseed rape plants were grown successively in pots of field soil for 4 weeks at a time, with six cycles of repeated vegetative planting in the same pot. Molecular analyses of the microbial communities after each cycle showed that the obligate parasite Olpidium brassicae infected the roots of oilseed rape within 4 weeks after the first planting (irrespective of the influence of rhizodeposits alone or in the presence of fresh detached root material), and consistently dominated the rhizosphere fungal community, ranging in relative abundance from 43 to 88 % when oilseed rape was grown more than once in the same soil. Fresh detached root material also led to a reduction in diversity within the soil fungal community, due to the increased relative abundance of O. brassicae. In addition, rhizosphere bacterial communities were found to have a reduced diversity over time when fresh root material was retained in the soil. In the second glasshouse experiment, the effect of incorporating mature, field-derived oilseed rape crop residues (shoots and root material) on microbial communities associated with subsequently grown oilseed rape was investigated. As before, molecular analyses revealed that O. brassicae dominated the rhizosphere fungal community, despite not being prevalent in either the residue material or soil fungal communities.     

Phosphorus nutrition of white lupine grown on a Calciaquoll

Abstract

White lupine (Lupinus albus L.) and navy bean (Phaseolus vulgaris L.) are legumes with and without the proteoid root characteristic, respectively. The influence of P (0, 40, 80, and 120 mg/kg of NaH₂PO₄‐P) and source of nitrogen (N) [inoculation with Bradyrhizobium lupini or Rhizobium phaseoli vs. NH₄NO₃] on phosphorus (P) uptake by navy bean and white lupine grown on a Calciaquoll low in available P and N was studied under greenhouse conditions. Both species responded to P fertilizer, but P accumulation in navy bean tops was much greater than in white lupine. Inoculated white lupine plants accumulated more P than their NH₄NO₃‐treated counterparts. Growth and P uptake of 17‐day‐old flax (Linum usitatissium L.) plants grown in pots previously planted to white lupine, but not in their navy bean counterparts, was severely retarded. White lupine did not enhance the availability of soil P in the calcareous soil.     

The effects of potassium fertilization on water‐use efficiency in crop plants

The supplies of water and nitrogen to a plant during its critical stages of growth are the main factors that define crop yield. A crop experiences irregular water deficits during its life cycle in rain‐fed agriculture. An effective anti‐stress‐oriented approach therefore ought to focus on increasing the units of water productivity. The main objective of this conceptual review is to confirm that adequate K management can be used as an important tool to alleviate the negative effects of water deficit on plant growth, yield‐component formation, and yield. The French and Schultz approach of using the water‐limited yield (WLY) was modified in this review into a graphical form and was used to discriminate between yield fractions that depended on the volume of transpired water from those that were induced by K fertilizer. By using this method, it was possible to demonstrate the extent of several crop (winter wheat, spring triticale, maize, sugar beet) responses to the K supply. Yield increases resulting from K application mostly appeared under conditions of mild water deficit. As described for sugar beet, finding the critical period of crop K sensitivity is a decisive step in understanding its impact on water‐use efficiency. It has been shown that an insufficient supply of K during crucial stages in the yield formation of cereals (wheat, spring triticale), maize, and sugar beet coincides with a depressed development in the yield components. The application of K fertilizer to plants is a simple agronomic practice used to increase crop tolerance to a temporary water shortage. It may be that the improvement of a plant's access to K during mild water‐deficiency stress will increase water uptake by the root cells, which in turn increases their osmotic potential and thereby allows extension growth. This growth in turn promotes access to other mineral elements (including nitrogen) and water, which favor plant growth and yield.     

Simulation of effects of soil bulk density and p addition on k uptake by soybeans

Abstract

Increasing soil bulk density has been shown to reduce root growth and decrease K uptake by soybeans (Glycine max L. Merrill). Changing soil bulk density also affects soil buffer power, b, and effective diffusion coefficient, De, which affect K influx. The relative decrease in K uptake due to reduced root growth as compared to reduced K influx is not known. Addition of P may affect root growth and P influx properties of plant roots. The objectives of this paper were (1) to use the Cushman mechanistic model to simulate the effect of changing soil bulk density and soil P on K uptake by soybeans, and (2) to determine the parameters that are changed by changes in bulk density and added P and their effect on K uptake. Plant and soil data of an experiment where Williams soybeans were grown for 21 days in pots of Raub (Aquic Argiudoll) silt loam with factorial treatments of two rates of K (0 and 100 mg K kg^‐1 soil), two rates of P (0 and 100 mg P kg^‐1 soil), and two bulk densities (1.25 and 1.45 g cm^‐3 ) were used to verify the model. Plant and soil parameters for the model were measured independently of the verification experiment. Predicted K (y) uptake agreed with observed uptake (x) (y = 1.09x‐0.19; r = 0.97) for the P x K factorial and (y = 1.19X‐0.22; r = 0.90) for the K x soil bulk density factorial treatments. In a sensitivity analysis, the model predicted a maximal K influx at a soil bulk density of 1.38 g cm^‐3. The greatest effect of soil bulk density on K uptake was due to reduction of root growth. Increased K uptake as a result of P addition was because of the effect on root growth.     

Phosphorus uptake from struvite is modulated by the nitrogen form applied

Next to nitrogen, phosphorus (P) is the most limiting nutrient for plant production worldwide. To secure food production, new nutrient management strategies using alternative P sources instead of mined P fertilizers need to be implemented. Struvite (MgNH₄PO₄ · 6 H₂O) is a promising example of a recycled mineral P fertilizer. Besides positive agronomic results regarding crop yields, further investigations are required to improve the use efficiency of the product and thereby increase its value. Using an automated plant phenotyping platform, we investigated the dynamic response to struvite by two plant species (lupine and maize) with diverse P acquisition strategies in an acidic sandy substrate. Although at three weeks after germination both maize and lupine had reduced leaf area in the struvite treatments compared to the commercial triple superphosphate (TSP), from week four onwards struvite plants grew larger than the TSP‐treated plants, indicating a slow release fertilizing effect. Greater P uptake efficiency (g / root length), but reduced root length were observed in the combined treatment of struvite and ammonium, in comparison to struvite and nitrate. We propose that rhizosphere acidification in response to ammonium uptake may enhance P recovery from struvite. A possible additional acidification effect by lupine root exudation might explain the higher P uptake efficiency in this species compared to maize. We conclude that struvite combined with ammonium can be used as a sustainable slow‐release P fertilizer on acidic sandy soils.     

Efficiency of Potassium Fertilization and Salicylic Acid on Yield and Nutrient Accumulation of Sugar Beet Grown on Saline Soil

Rising soil salinity has been a major problem in the soils of Egypt in recent decades. Potassium fertilization and salicylic acid (SA) play an important role in promoting plants to tolerate salt stress and increased the yield of sugar beet crop. A field experiment on sugar beet (Beta vulgaris L.) grown on saline soil was carried out during 2014 growing season in Port Said Governorate, Egypt, to study the effect of potassium fertilization of the soil at applications of 0, 100, 150, and 200 kg potassium (K) ha^?1 and foliar spray of SA by solution of 1000 mg L^?1, twice (1200 L ha^?1 each time) on yield and nutrient uptake. Application of 200 kg K ha^?1 in combination with salicylic foliar spray gave the highest root length, root diameter, shoot and root yield, sucrose, juice purity percentage, gross sugar yield, and white possible extractable sugar, nitrogen (N), phosphorus (P), and potassium (K) content, and uptake of sugar beet. The highest increase in sucrose (20%) as well as white possible extractable sugar (184%) was obtained by 200 kg K ha^?1 in combination with salicylic foliar spray compared with untreated soil with potassium fertilization and without salicylic foliar spray.     

Utilization of organic phosphorus sources by oilseed rape,sunflower, and soybean

We evaluated the ability of Brassica napus L. (oilseed rape), Helianthus annus L. (sunflower), and Glycine max L. (soybean) plants grown inoculated with or without bacteria to utilize organic P sources. Plants were supplied with inorganic (dibasic sodium phosphate) and organic P sources (phytate and glucose phosphate) at three concentrations and grown for 40 d under sterile conditions. Three inoculation treatments were compared: control (non‐inoculated plants), inoculation with Bacillus amyloliquefaciens BNM340, and inoculation with Pseudomonas fluorescens BNM296 (two bacteria with proven phytase activity). Oilseed rape, sunflower and soybean could utilize organic P sources. For example, when phytate (0.5 mM) P was used as the external P source, the increase factors over the no‐P treatments were 4.5, 1.4, and 1.4 for oilseed rape, sunflower, and soybean P uptake, respectively. When glucose 1‐phosphate disodium salt (G1P, 0.5 mM) was the P source, the increase factors were 8.8, 1.7, and 1.9 respectively. Positive responses to the organic P sources were found for the biomass accumulation of oilseed rape and soybean but not for sunflower. The inoculation with bacteria did not exert a promoting effect on P uptake. We demonstrate that the three species can effectively use organic P sources. The existence of crop plants that are more efficient in the utilization of different soil P sources would be particularly beneficial to improve P recycling and use of P fertilizers in agriculture.     

Effect of supplying P to a portion of the soybean root system on root growth and P uptake kinetics 1

Abstract

Knowledge of the effect of supplying P to portions of the soybean (Glycine max L. Merr) root system on P influx kinetics and root growth is important in developing P fertilizer placement practices for efficient fertilizer use. The objective of this research was to determine the effect of restricting P supply to portions of the root system on plant P status, root growth, and P influx kinetics. Two solution experiments were conducted in a controlled climate chamber. Phosphorus influx kinetics were determined on 25‐day‐old soybean plants that had been grown with 100, 75, 50, 25, and 12.5% of their roots initially exposed to P. Phosphorus influx kinetics were also measured on 25‐day‐old plants that had been P‐starved for the last 1, 2, 4, and 6 days prior to the determining P influx kinetics in order to relate plant P status to P influx kinetics.

Reducing the portion of the roots supplied with P reduced P uptake. This resulted in a reduction in plant P concentration and was related to a 3.41‐fold increase in maximum P influx measured on 25‐day‐old plants. Restricting the proportion of roots supplied with P had no significant effects on the Michaelis‐Menten constant or on the concentration in solution where net influx was zero. Root growth rate of the roots in the P containing solution was not significantly different from those in the ‐P solution.

Phosphorus uptake was correlated with final root surface area exposed to P (r² = 0.88??). Starving the plants for P reduced P concentration in the shoot and root and this resulted in as much as a 1.68‐fold increase in maximum influx.     

The effect of co‐composted cabbage and ground phosphate rock on the early growth and P uptake of oilseed rape and perennial ryegrass

Phosphorus (P) availability to crops in organic systems can be a major issue, with the use of readily available forms often restricted. One product that can be used in organically managed systems, that is also relatively easily accessible to growers, is phosphate rock, although its solubility and therefore crop availability is often poor. One possible approach to improve this situation is co‐composting phosphate rock with selected organic waste materials. Various ratios of phosphate rock and cabbage (Brassica oleracea L.) residues were co‐composted and the products tested at different rates of application. The effects were assessed over 12 weeks using oilseed rape (Brassica napus L.) and perennial ryegrass (Lolium perenne L.) as bioassay crops in a pot experiment. At harvest, estimates of P derived from cabbage and phosphate rock for the lowest of two rates of compost were ≈ 2 and 10 mg P pot^–1 for oilseed rape, compared to 5 and 2 mg P pot^–1 for perennial ryegrass, respectively. Roots tended to have higher P concentrations than shoots. The crops showed differences in their abilities to access various P sources, with oilseed rape effectively taking P from phosphate rock, whereas perennial ryegrass was more effective at accessing cabbage‐derived P (the main substrate in the compost). Oilseed rape was able to take up 20% of the total P applied as phosphate rock, whereas perennial ryegrass took up less than 5% of the total P applied from this material. Both pre‐ and post‐application solubilisation/transformation mechanisms were involved in supplying plant‐available P. Quantifying the relative contribution from individual P sources remains problematic even within this relatively simple system.     

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.     

Fermentation of sugar beet waste by Aspergillus niger facilitates growth and P uptake of external mycelium of mixed populations of arbuscular mycorrhizal fungi

Sugar beet waste has potential value as a soil amendment and this work studied whether fermentation of the waste by Aspergillus niger would influence the growth and P uptake of arbuscular mycorrhizal (AM) fungi. Plants were grown in compartmentalised growth units, each with a root compartment (RC) and two lateral root-free compartments (RFC). One RFC contained untreated soil while the other RFC contained soil, which was uniformly mixed with sugar beet waste, either untreated (SB) or degraded by A. niger (ASB) in a rock phosphate (RP)-supplied medium. The soil in each pair of RFC was labelled with ³³P and ³²P in order to measure P uptake by the AM fungal mycelium, of which length density was also measured. Whole cell fatty acid (WCFA) signatures were used as biomarkers of the AM fungal mycelium and other soil microorganisms. The amount of biomarkers of saprotrophic fungi and both Gram-positive and Gram-negative bacteria was higher in SB than in ASB treatments. Whilst ASB increased growth and activity of AM mycelium, SB had the opposite effect. Moreover, shoot P content was increased by the addition of ASB, and by inoculation with AM fungi. Modification of soil microbial structure and production of exudates by A. niger, as a consequence of fermentation process of sugar beet waste, could possibly explain the increase of AM growth in ASB treatments. On the other hand, the highest P uptake was a result of the solubilisation of rock phosphate by A. niger during the fermentation.     

Effect of organic amendments on uptake of selenium and biochemical grain composition of wheat and rape grown on seleniferous soils in northwestern India

Crops grown in seleniferous soil may accumulate selenium (Se) to levels considered highly toxic for animal and human consumption. Furthermore, higher Se content in plant tissues leads to considerable deterioration in product quality. Application of organic amendments plays an important role in improving soil physical, chemical, and biological conditions and influencing nutrient availability. A field trial was conducted to evaluate the effect of organic amendments, namely poultry manure (PM), sugar cane press mud (SCPM), and farmyard manure (FYM), on Se uptake and grain quality of wheat and oilseed rape grown on a seleniferous soil in Punjab, India. Selenium accumulation by wheat and oilseed rape grains decreased significantly (75%–95%) with the application of PM and SCPM, while FYM application resulted in a significant decrease (23%) only in case of wheat grains. The amount of Se associated with seed proteins varied in proportion to its uptake under different treatments. Quality of wheat grains improved considerably with respect to total soluble sugars, reducing sugars, starch, lipids, and sulfur concentrations only after application of SCPM and PM. Treating a Se‐contaminated soil with organic amendments significantly increased the oil concentration and changed the proportion of various fatty acids in rape grains. It is concluded that applying organic amendments to Se‐contaminated soils can alleviate the deleterious effects of Se and restore the nutritional quality of grains.     

Interactive effects of plant growth–promoting rhizobacteria and organic fertilization on P nutrition of Zea mays L. and Brassica napus L.

The organic matter supply can promote the dispersal and activity of applied plant growth–promoting rhizobacteria (PGPR), but the complementary effect of organic fertilization and PGPR application on the turnover of P is scarcely known. The effects of the application of two PGPR strains (Pseudomonas fluorescens strain DR54 and Enterobacter radicincitans sp. nov. strain DSM 16656) alone and in combination with organic fertilization (cattle manure and biowaste compost) on growth and P uptake of maize (Zea mays L.) and oilseed rape (Brassica napus L.) were investigated under semi–field conditions. Furthermore, P pools and phosphatase activities in soil and the arbuscular mycorrhizal colonization of maize were examined. The organic‐fertilizer amendments increased the growth and P uptake of both plant species and the soil P pools. The application of the E. radicincitans strain increased P uptake of oilseed rape when no organic fertilizer was added. Furthermore, the application of both bacterial strains increased the activities of phosphatases under both plant species. Here, the effect of the PGPR application even exceeded the effect of organic fertilization. The magnitude of this effect varied between the different fertilizing treatments and between the two bacterial strains. Phosphatase activities were increased to the greatest extent after application of P. fluorescens in the unfertilized soil. Under rape increases of 52% for acid phosphatase activities (ACP), 103% for alkaline phosphatase activities (ALP), and 133% for phosphodiesterase (PDE) were observed therewith. In the unfertilized soil, the application of P. fluorescens also resulted in a strong increase of the arbuscular mycorrhizal colonization of maize. We conclude that application of PGPR can promote the P mobilization and supply of crops in P‐deficient soils, however, in combination with organic fertilization these effects might be masked by a general improved P supply of the crops. Interactive effects of applied bacterial strains and organic fertilization depend on the sort of organic fertilizer and crop species used.     

Effect of Root Zone Temperature on Oilseed Rape (Brassica napus) Response to Boron

Abstract

Oilseed rape (Brassica napus) is sensitive to low boron (B) supply, and its growth response to B may be influenced by soil temperature. To test the relationship between B and temperature, oilseed rape (cv. Hyola 42) seedlings were grown at 10°C (low) root zone temperature (RZT) with B supply from deficient to adequate B levels until growth of low B plants just began to slow down. Half of the pots were then transferred to 20°C (warm) RZT for 11 days before they were moved back to 10°C RZT for the final 4 days. Both plant dry mass and B uptake increased after plants were exposed to warm RZT. However, plant B deficiency was exacerbated by warm RZT in low B plants because of increased relative growth rate and shoot–root ratio without a commensurate increase in B uptake rate. It is concluded that RZT above the critical threshold for chilling injury in oilseed rape can nevertheless affect the incidence of B deficiency by altering shoot–root ratio and hence the balance between shoot B demand and B uptake.     

Identification of Phosphorous Efficient Germplasm in Oilseed Rape

ABSTRACT

Plant species and genotypes within one species may significantly differ in phosphorus (P) uptake and utilization when they suffer from P starvation. The objective of this research was to screen P-efficient germplasm of oilseed rape (Brassica napus L.) and analyze the possible mechanism responsible for P efficiency by two-steps screening experiments and validation of P efficiency. Phosphorus efficiency coefficient at seedling stage, namely, ratio of shoot dry weight under low P to that under adequate P (PECS) of 194 oilseed rape cultivars varied from 0.050 to 0.62 and was significantly related with shoot dry weight under low P level (r = 0.859??, P < 0.01). Oilseed rape cultivar ‘Eyou Changjia’ presented the highest P efficiency coefficient in each growth stage and had the highest seed yield at low P, whereas oilseed rape cultivar ‘B104-2’ was the most sensitive to low P stress among the 12 candidate cultivars obtained from the two-steps screening experiments. Under low P condition in validation experiments of soil and solution cultures, ‘Eyou Changjia’ could produce much more dry matter and acquire more P than ‘B104-2.’ Moreover, P efficient coefficient obtained from the pot experiment was comparable to those from the field experiment. This might be attributed to high P uptake efficiency for ‘Eyou Changjia’ when it suffered from low-P stress. Comparison of results from the hydroponics with those from the pot and field experiments led to the conclusion that the P uptake efficiency in the hydroponics is highly related to that in soil culture conditions. These results show that there are large genotypic differences in response to phosphorus deficiency in oilseed rape germplasm (Brassica napus L.) and ‘Eyou Changjia’ is P-efficient and ‘B104-2’ is P-inefficient. By comparing these results further, the mechanism responsible for P efficiency was suggested to be mainly due to high P uptake efficiency by forming larger root system, and improving the ability of mobilizing and acquiring soil P in P-efficient oilseed rape under the condition of P starvation.     

Ability of various water‐insoluble fertilizers to supply available phosphorus in hydroponics to plant species with diverse phosphorus‐acquisition efficiency: Involvement of organic acid accumulation in plant tissues and root exudates

Previous studies describe the suitability of a new type of phosphorus (P) fertilizer, called “rhizosphere‐controlled fertilizer” (RCF), to supply available P to plants while reducing soil phosphorus fixation. In order to explore the involvement of organic acid root exudation in P uptake from RCF, we investigated the relationship between shoot and root P concentrations, and the concentration of the main polycarboxylic organic acids in roots, shoots, and plant exudates. Plant species with different P‐acquisition efficiency (low: maize; medium: chickpea; high: lupin) were grown in hydroponics with three different P fertilizers: The water‐insoluble P fraction of RCF (RCF); Phospal, a slow‐release source of phosphate composed of calcium and aluminum phosphates (PH); monopotassiumphosphate (KP), and a control treatment without P (P–). RCF was as efficient as KP in supplying P to plants in the case of chickpea and lupin, and slightly less efficient than KP in maize. However, P from PH was not available for maize and less available compared to KP and RCF in chickpea and lupin. This variation reflects the different efficiencies in P acquisition for the three plant species. Except in the case of maize, plants receiving KP presented the lowest concentration of organic acids in roots and exudates, while those plants suffering severe P deficiency (P– and PH) showed the highest organic acid concentration. However, RCF had a high concentration of organic acids in roots and exudates, as well as a high P concentration in the shoot indicating that P uptake from RCF is enhanced due to root release and action of specific organic acids.     

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.     

A conceptual framework and an empirical test of complementarity and facilitation with respect to phosphorous uptake by plant species mixtures

Plant species have different traits for mobilizing sparingly soluble phosphorus (P) resources, which could potentially lead to overyielding in P uptake by plant species mixtures compared to monocultures due to higher P uptake as a result of resource (P) partitioning and facilitation. However, there is circumstantial evidence at best for overyielding as a result of these mechanisms. Overyielding (the outcome) is easily confused with underlying mechanisms because of unclear definitions. We aimed to define a conceptual framework to separate outcome from underlying mechanisms and test it for facilitation and complementarity with respect to P acquisition by three plant species combinations grown on four soils. Our conceptual framework describes both mechanisms of complementarity and facilitation and outcomes (overyielding of mixtures or no overyielding) depending on the competitive ability of the species to uptake the mobilized P. Millet/chickpea mixtures were grown in pots on two calcareous soils mixed with calcium-bound P (CaP) and phytate P (PhyP). Cabbage/faba bean mixtures were grown on both acid and neutral soils mixed with P-coated iron (hydr)oxide (FeP) and PhyP. Wheat/maize mixtures were grown on all four soils. Rhizosphere carboxylate concentration and acid phosphatase activity (mechanisms) as well as plant P uptake and biomass (outcome) were determined for monocultures rhizosphere and species mixtures. Facilitation of P uptake occurred in millet/chickpea mixtures on one calcareous soil. We found no indications for P acquisition from different P sources, neither in millet/chickpea, nor in cabbage/faba bean mixtures. Cabbage and faba bean on the neutral soil differed in rhizosphere acid phosphatase activity and carboxylate concentration, but showed no overyielding. Wheat and maize, with similar root exudates, showed overyielding (the observed P uptake being 22% higher than the expected P uptake) on one calcareous soil. We concluded that although differences in plant physiological traits (root exudates) provide necessary conditions for complementarity and facilitation with respect to P uptake from different P sources, they do not necessarily result in increased P uptake by species mixtures, because of the relative competitive ability of the mixed species.