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.     

Root-induced changes in the rhizosphere: Importance for the mineral nutrition of plants

Root-induced changes in the rhizosphere may affect mineral nutrition of plants in various ways. Examples for this are changes in rhizosphere pH in response to the source of nitrogen (NH₄-N versus NO₃-N), and iron and phosphorus deficiency. These pH changes can readily be demonstrated by infiltration of the soil with agar containing a pH indicator. The rhizosphere pH may be as much as 2 units higher or lower than the pH of the bulk soil. Also along the roots distinct differences in rhizosphere pH exist. In response to iron deficiency most plant species in their apical root zones increase the rate of H⁺ net excretion (acidification), the reducing capacity, the rate of Fe^III reduction and iron uptake. Also manganese reduction and uptake is increased several-fold, leading to high manganese concentrations in iron deficient plants. Low-molecular-weight root exudates may enhance mobilization of mineral nutrients in the rhizosphere. In response to iron deficiency, roots of grass species release non-proteinogenic amino acids (?phytosiderophores”?) which dissolve inorganic iron compounds by chelation of Fe^III and also mediate the plasma membrane transport of this chelated iron into the roots. A particular mechanism of mobilization of phosphorus in the rhizosphere exists in white lupin (Lupinus albus L.). In this species, phosphorus deficiency induces the formation of so-called proteoid roots. In these root zones sparingly soluble iron and aluminium phosphates are mobilized by the exudation of chelating substances (probably citrate), net excretion of H⁺ and increase in the reducing capacity. In mixed culture with white lupin, phosphorus uptake per unit root length of wheat (Triticum aestivum L.) plants from a soil low in available P is increased, indicating that wheat can take up phosphorus mobilized in the proteoid root zones of lupin. At the rhizoplane and in the root (root homogenates) of several plant species grown in different soils, of the total number of bacteria less than 1 % are N₂-fixing (diazotrophe) bacteria, mainly Enterobacter and Klebsiella. The proportion of the diazotroph bacteria is higher in the rhizosphere soil. This discrimination of diazotroph bacteria in the rhizosphere is increased with foliar application of combined nitrogen. Inoculation with the diazotroph bacteria Azospirillum increases root length and enhances formation of lateral roots and root hairs similarly as does application of auxin (IAA). Thus rhizosphere bacteria such as Azospirillum may affect mineral nutrition and plant growth indirectly rather than by supply of nitrogen.     

Crop species differ in root plasticity response to localised P supply

The effect of localised phosphorus (P) fertiliser placement and in particular, deep P fertiliser placement, on the comparative root growth and P uptake of fibrous vs tap‐rooted crops is not known. In this study, we examined the root growth and P uptake of wheat (Triticum aestivum L.), canola (Brassica napus L.), and narrow‐leaf lupin (Lupinus angustifolius L.) in a split‐root system and in columns with deep (19 cm) or shallow (5 cm) P fertiliser sources in glasshouse conditions. In the split‐root system, plants of all three species grown under heterogeneous soil P conditions absorbed more P and produced greater root and shoot biomass than those under homogeneous P supply. Root plasticity differed between species under heterogeneous soil P supply: canola and wheat allocated relatively more root biomass and root length to the high P zone than narrow‐leaf lupin. In the column experiment, there was no difference in the amount of P accumulated in shoots of any crops grown in the deep vs shallow P fertiliser treatments. Root proliferation occurred within the shallow and deep‐P fertiliser bands in all three species; however, root distribution above or below the bands did not differ between deep or shallow P fertiliser treatments in any species. Whilst root plasticity responses to heterogeneous soil P supply differed among species, root architecture (fibrous vs taproot) did not confer any advantage or disadvantage to the acquisition of P from deep vs shallow P fertiliser bands. Moreover, whilst roots proliferate in the vicinity of P fertiliser bands, root distribution outside of the bands appears to remain unaltered in both fibrous and tap‐rooted crops during early growth.     

Cluster Root Formation by Lupinus Albus is Modified by Stratified Application of Phosphorus in a Split-Root System

ABSTRACT

Cluster root formation by white lupin (Lupinus albus L. cv. Kiev Mutant) in response to stratified application of hydroxyapatite was examined in a split-root system. The system consisted of two vertical compartments, each divided horizontally into five 60-mm layers. Hydroxyapatite was applied to different layers at 150 mg phosphorus(P) kg^?1 soil. The proportion of dry biomass of cluster roots in the whole root system was significantly reduced when P concentration was high in shoots due to P application, suggesting that cluster root formation was regulated by the shoot P status. However, the cluster root percentage increased in the soil layer supplemented with P, and decreased in other layers, especially when P was applied in a deep layer. The formation of cluster roots is regulated by internal plant P status, but is also greatly affected by localized P supply. Heterogeneous P supply can modify the distribution of cluster roots.     

Cluster root allocation of white lupin (Lupinus albus L.) in soil with heterogeneous phosphorus and water distribution

Cluster roots are structures formed by many plants adapted to phosphorus (P)-deficient soils. We investigated the combined influence of spatial heterogeneity in soil water and P distribution on the allocation of cluster root formation in white lupin (Lupinus albus L.). In this study, single plants were grown at a low or a high rate of water supply in containers filled with a P-poor sand to which either no P was added or which was fertilized homogeneously or heterogeneously. Furthermore, heterogeneous soil water distribution was established in half of the containers by using a finer instead of a coarser sand in a lateral third of the containers. Plant growth increased with water supply rate, but P fertilization had no influence on shoot biomass production. Although overall cluster root production decreased with increasing homogeneous P supply, localized P fertilization had no effect on cluster root allocation. However, cluster roots were preferentially allocated in the soil sections with lower water availability when overall water supply rate was low. The results suggest that overall cluster root production was a systemic response to initial plant P status, while cluster root growth was stimulated locally in drier patches when overall water supply was limiting plant growth.     

Aluminium-Toleranz von Ackerbohne (Vicia faba), Lupine (Lupinus luteus), Gerste (Hordeum vulgare) und Roggen (Secale cereale). II. Mineralstoffgehalte in Sproß und Wurzeln in Abhängigkeit vom Aluminium-Angebot

Al tolerance of horse bean, yellow lupin, barley and rye. II. Mineral element concentrations in shoots and roots as affected by Al supply Inhibition of seminal root elongation by Al in solution culture gave the following ranking for Al tolerance: yellow lupin (Lupinus luteus ?Schwako”?) ? rye (Secale cereale ?Kustro)”? « horse bean (Vicia faba ?Herz Freya”?) > barley (Hordeum vulgare ?Roland”?). Exclusion from uptake by inactivation of Al outside the root was not responsible for the higher Al tolerance of lupin and rye, because comparable inhibition of root elongation occured at much higher Al concentration of the root and the root tips (5 mm) compared to barley and horse bean. The plant species differed considerable in nutrient concentrations of the roots: higher Ca concentrations in horse bean and rye, higher Mg concentrations in rye and lupin and higher P concentration in lupin. Al supply reduced Ca and Mg concentrations (Ca > Mg) in shoots and roots of all species. P concentrations were hardly affected. The nutrient concentrations in the root tips did not indicate that induction of nutrient deficiency was responsible for the effect of Al on root elongation and Al sensitivity of barley and horse bean. The considerable differences in Ca, Mg and P concentrations of the roots between the Al-tolerant plant species rye and lupin do not suggest a common physiological mechanism responsible for Al tolerance.     

Root exudation of sugars,amino acids,and organic acids by maize as affected by nitrogen,phosphorus, potassium,and iron deficiency

Root exudates play a major role in the mobilization of sparingly soluble nutrients in the rhizosphere. Since the amount and composition of major metabolites in root exudates from one plant species have not yet been systematically compared under different nutrient deficiencies, relations between exudation patterns and the type of nutrient being deficient remain poorly understood. Comparing root exudates from axenically grown maize plants exposed to N, K, P, or Fe deficiency showed a higher release of glutamate, glucose, ribitol, and citrate from Fe‐deficient plants, while P deficiency stimulated the release of γ‐aminobutyric acid and carbohydrates. Potassium‐starved plants released less sugars, in particular glycerol, ribitol, fructose, and maltose, while under N deficiency lower amounts of amino acids were found in root exudates. Principal‐component analysis revealed a clear separation in the variation of the root‐exudate composition between Fe or P deficiency versus N or K deficiency in the first principal component, which explained 46% of the variation in the data. In addition, a negative correlation was found between the amounts of sugars, organic and amino acids released under deficiency of a certain nutrient and the diffusion coefficient of the respective nutrient in soils. We thus hypothesize that the release of dominant root exudates such as sugars, amino acids, and organic acids by roots may reflect an ancient strategy to cope with limiting nutrient supply.     

Exudation of organic anions by roots of cabbage,carrot, and potato as influenced by environmental factors and plant age

A previous study demonstrated that cabbage was P efficient compared to carrot and potato. However, calculating plant P uptake by a mechanistic simulation model based on P transport by diffusion and mass flow, P uptake of roots according to the Michaelis‐Menten kinetics, and morphological root characteristics including root hairs, revealed that these parameters could explain only 2/5 of the total P uptake of cabbage, but 4/5 of that of carrot and potato (Dechassa et al., 2003). Therefore, it was hypothesized that a higher root exudation of organic anions may enhance P mobilization and hence P uptake of cabbage. The objective of this research was to determine root exudation of organic anions by the three species, and to investigate the influence of plant age and dark/light period on organic‐anion exudation by cabbage. Experiments were conducted in a growth chamber in nutrient solution with or without P. Organic anions were determined in root exudates and in root tissue. With cabbage and potato, P deficiency induced exudation of citrate and succinate, respectively. Citrate‐exudation rate of P‐deficient cabbage plants was correlated with accumulation of citrate in root tissue. In contrast, high succinate‐exudation rates in potato were not correlated with an increased concentration in root tissue. For carrot, no change was observed in the exudation of any of the organic anions in response to P deficiency. The results also showed that succinate‐ and citrate‐exudation rates of cabbage roots increased with increased plant age. There was also a significant increase in exudation rates of organic anions of cabbage roots during the light period of the day. It was concluded that cabbage had the ability to exude large amounts of citrate in response to P deficiency by which it can additionally enhance its P‐uptake efficiency, whereas carrot and potato showed little evidence of possessing such a mechanism.     

Subsoil rhizosphere modification by chickpea under a dry topsoil: implications for phosphorus acquisition

Chickpea (Cicer arietinum L.) roots exude carboxylates. While chickpea commonly grows where the topsoil dries out during crop growth, the importance of carboxylate exudation by the roots and mobilization of soil P from below the dry topsoil has not been examined. The study investigates the response of carboxylate exudation and soil P mobilization by this crop to subsoil P fertilizer rate. In constructed soil columns in the glasshouse, the P levels (high, low, and nil P) were varied in the well‐watered subsoil (10–30 cm), while a low level of P in the dry topsoil (0–10 cm) was maintained. At flowering, rhizosphere carboxylates and rhizosphere soil from topsoil and subsoil roots were collected separately and analyzed. The concentration of total carboxylates per unit rhizosphere mass in the subsoil was nearly double that of the topsoil. Plants depleted sparingly soluble inorganic P (P_i), NaOH‐P_i, and HCl‐P_i, along with the labile P_i (water soluble and NaHCO₃‐Pi). The P depletion by plants was greater from the subsoil than the topsoil. The study concluded that depletion of sparingly soluble P from the chickpea rhizosphere in the subsoil was linked with the greater levels of carboxylates in the rhizosphere. These findings indicate that chickpea, with its deep rooting pattern, can increase its access to subsoil P when the topsoil dries out during crop growth by subsoil rhizosphere modification.     

VARIATION IN PHOSPHORUS EFFICIENCY AMONG BRASSICA CULTIVARS II: CHANGES IN ROOT MORPHOLOGY AND CARBOXYLATE EXUDATION

Increased phosphorus (P) efficiency is needed to sustain agriculture productivity on soils with low available P. Significant differences were found among Brassica cultivars for growth, P utilization, and remobilization under P deficiency (see our companion paper, Aziz et al., 2011a). To identify the possible mechanisms of P acquisition from low soluble P compounds, four cultivars (‘Rainbow’, ‘CON-1’, ‘Dunkeld’, and ‘Peela Raya’) were selected to ascertain the relationship of their differential P acquisition and growth with their root length in soil and with organic acid release pattern in solution culture experiments. For this purpose their growth and P acquisition from phosphate rock (PR) was compared with calcium di-hydrogen phosphate (Ca-P) when adding uniform dose of 100 mg P kg^?1 soil separately from the two sources. Biomass accumulation, root length, root fineness, plant P uptake and ash alkalinity was significantly (P < 0.01) different in plants of all the four cultivars when supplied with PR or Ca-P in soil. Minimum biomass produced by ‘Peela Raya’ grown with either P source was followed by ‘CON-1’, ‘Dunkeld’, and ‘Rainbow’ in ascending order. Shoot dry matter production had a significant positive correlation with root dry matter production (r = 0.85, P < 0.01), root length (r = 0.59, P < 0.05) and root P uptake (r = 0.95, P < 0.01). Cultivars varied significantly for organic acid secretion in solution culture experiment. Higher quantities of secreted citric acid, malic acid and tartaric acid in solution culture experiment were measured for ‘Rainbow’ and ‘Dunkeld’ cultivars. Efficient performance of these two cultivars for growth and P uptake was associated with their longer roots and more secretion of organic acids especially citric acid.     

Changes in net proton release by roots of intact maize plants after local nutrient supply in a split root system

The effect of local nutrient supply to maize roots (Zea mays L. cv. Blizzard) on net proton release was studied using the split root technique (SRNS, SRCa) to compare plants that were cultivated with their roots completely in either nutrient solution (NS) or 0.1 mM CaSO₄ (Ca). Roots in NS released more protons than roots in Ca. This higher net proton release was associated with significantly higher ATP concentrations in the root tissue. Higher net proton release and ATP concentrations were also observed after a 4 h lag phase when 20 μM abscisic acid were exogenously applied to roots in 0.1 mM CaSO₄. It is suggested that higher metabolic activity in roots supplied with nutrients increased ATP concentrations and thus the substrate supply of the plasma membrane H⁺ ATPase. When only half of the root system was supplied with nutrient solution with the other half bathed in 0.1 mM CaSO₄, the roots in the SRNS compartment released significantly higher amounts of protons relative to the NS control plants. Conversely, roots in the SRCa compartment showed net proton uptake in contrast to the roots of control plants in 0.1 mM CaSO₄ which significantly acidified the root medium. These differences in proton release by roots in the split root system and control roots could not be explained in terms of differences in ATP concentrations. It is therefore suggested that an internal signal may lead to a modification of the plasma membrane H⁺ ATPase as shown earlier during plant adaptation to low pH in the root medium.     

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.     

Phosphorus uptake and rhizosphere properties of intercropped and monocropped maize, faba bean, and white lupin in acidic soil

Little information is available on phosphorus (P) uptake and rhizosphere processes in maize (Zea mays L.), faba bean (Vicia faba L.), and white lupin (Lupinus albus L.) when intercropped or grown alone in acidic soil. We studied P uptake and soil pH, carboxylate concentration, and microbial community structure in the rhizosphere of maize, faba bean, and white lupin in an acidic soil with 0–250 mg P (kg⁻¹ soil) as KH₂PO₄ (KP) or FePO₄ (FeP) with species grown alone or intercropped. All plant species increased the pH compared to unplanted control, particularly faba bean. High KP supply (>100 mg P kg⁻¹) significantly increased carboxylate concentration in the rhizosphere of maize. The carboxylate composition of the rhizosphere soil of maize and white lupin was significantly affected by P form (KP or FeP), whereas, this was not the case for faba bean. In maize, the carboxylate composition of the rhizosphere soil differed significantly between intercropping and monocropping. Yield and P uptake were similar in monocropping and intercropping. Monocropped faba bean had a greater concentration of phospholipid fatty acids in the rhizosphere than that in intercropping. Intercropping changed the microbial community structure in faba bean but not in the other corps. The results show that P supply and P form, as well as intercropping can affect carboxylate concentration and microbial community composition in the rhizosphere, but that the effect is plant species-specific. In contrast to previous studies in alkaline soils, intercropping of maize with legumes did not result in increased maize growth suggesting that the legumes did not increase P availability to maize in this acidic soil.     

Plant growth,leaf photosynthesis,and nutrient‐use efficiency of citrus rootstocks decrease with phosphite supply

Some formulations of phosphite (Phi) have been recommended as a source of P nutrition for several crops including citrus even though there are known negative effects of Phi on plant growth. Changes in plant growth and metabolism after Phi application should be reflected in altered nutrient‐use efficiency and leaf photosynthesis. We carried out a greenhouse study using seedlings of two contrasting citrus (Citrus spp.) rootstocks, Carrizo citrange (CC) and Smooth Flat Seville (SFS), growing in either aerated hydroponic culture or sterilized native sandy soil. Plants were subjected to four P treatments: No P (control, P₀); 0.5 mM P_i (PO₄‐P); 0.25 mM P_i + 0.25 mM Phi (P_i + Phi), or 0.5 mM Phi (Phi). Photosynthetic characteristics, concentrations of total P (P_t) and soluble PO₄‐P or PO₃‐P in leaves and roots, and plant growth were evaluated after 80–83 d P treatments. Overall, the P_i plants had the highest P_t (total P) and total plant dry weight while the P₀ plants had the lowest P_t but highest total root length and root‐to‐shoot ratio. Leaf chlorophyll (SPAD readings) and net assimilation of CO₂ (A_CO2) of the P₀ and Phi plants were similarly lower than those of P_i and P_i + Phi plants. Growth responses of the P_i + Phi treatment were intermediate between the P_i and Phi treatments. Although Phi increased P_t and soluble‐PO₄‐P concentration in leaves and roots above the P₀ treatment, this did not translate into increased plant growth. In fact, the Phi treatment had some phytotoxic symptoms, impaired P‐ and N‐utilization efficiency for biomass production as well as lower nutrient‐use efficiency in the photosynthetic process. Thus, these two rootstocks could not use Phi as a nutritional source of P.     

旱后复水条件下磷添加对白羊草根系形态及氮磷累积的影响

植物生长及养分利用特征可揭示半干旱区植物对多变水肥环境条件的适应策略。在白羊草分蘖期设置2个供水条件(正常供水和干旱胁迫21天后复水)和2个磷添加水平(复水当日1 kg干土添加0,0.2 g P_2O_5),2周后测定其根冠生物量、根系形态以及氮磷含量。结果表明,旱后复水条件下,磷添加后白羊草根冠生物量、总生物量和根冠比无显著变化,总根长和根表面积显著增加27.1%和24.1%,比根长和比根面积分别显著增加18.3%和15.9%,根系平均直径显著降低1.3%;白羊草地上部、根系和整株磷含量分别显著增加61.1%,35.8%和49.6%,磷累积量分别显著增加68.6%,52.0%和61.3%,氮磷比显著降低。除地上部氮累积量外,各水分和磷处理下白羊草地上部、根系和整株氮磷累积量与总根长和根表面积呈显著正相关关系。本研究表明,根长和根表面积增加是白羊草响应水肥环境条件改善的主要策略。     

玉米苗期根系生长与耐低磷的关系

在田间筛选试验的基础上,利用两个磷高效(181和186)、两个磷低效(153和197)玉米自交系,进一步研究了这些自交系苗期耐低磷能力差异及其与根系生长的关系。结果表明,在低磷胁迫(P.5.78.mg/kg)下,所有自交系玉米地上部重量、初生根重、次生根重及磷累积量降低,但磷高效自交系181和186受影响程度显著小于153和197。在试验所处的玉米生育时期(6叶龄),磷对所用自交系的初生根及次生根数量没有影响。比较181和197的根系形态,在低磷胁迫下,磷低效自交系197的初生根侧根长、轴根长均显著下降,磷高效自交系181则下降幅度很小。而且,低磷使181初生根的侧根/轴根比值、根长度/根重比值较高。说明低磷胁迫下,181自交系可以将根中的有限的养分及干物质作更合理的分配,促进细根的生长,从而获得较长的根系。     

不同供磷状况下CO2浓度升高对番茄根系生长及养分吸收的影响

采用培养试验研究了磷缺乏与正常供磷条件下,CO₂浓度由350μL/L升高至800μL/L苗期番茄的生物量、根系特征和不同器官N、P、K养分含量的变化。结果表明,无论缺磷与否,CO₂浓度升高均能显著增加番茄地上部及根系的干物质积累量,提高根冠比。在磷缺乏条件下,CO₂浓度升高对番茄根系生长的促进主要表现为增加根系的体积和表面积;而在磷正常供应条件下主要表现为同时增加根体积和分根数,有利于形成强壮的根系。在两种供磷水平下,CO₂浓度升高对番茄各器官的N、P、K含量产生不同的稀释效应,但N、P、K总积累量却随CO₂浓度升高而显著增加;而且CO₂浓度与供P水平对番茄植株的N、P、K积累量具有极显著的正交互效应。     

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.     

玉米细根形成和土壤中无机磷分级平衡对缺磷的特异性响应

Plants have diverse strategies to cope with phosphorus (P) deficiency. To better understand how maize responds to P deficiency, a field experiment with two P levels, 0 and 100 kg P2O5 ha-1 (P0 and P100, respectively), was carried out as a part of a long-term Pfertilizer field trial. Plant and soil analyses showed that P-deficient maize reduced its growth rate, increased P use efficiency, and formed more thin roots with the diameter less than 0.6 mm at jointing and silking stages, compared to the plants treated with P100. Further, there were no differences in major inorganic P fractions (Ca 2 -P, Ca 8 -P, Al-P, Fe-P, occluded P and Ca 10 -P) between the rhizospheric and bulk soils at each harvest, even when soil Olsen-P was only 1.38 mg kg-1 . These results suggested that maize responded to P deficiency by reducing the internal P demand for growth and increasing P acquisition ability by favorable root morphological alteration at low carbon cost.