Carbon Turnover in the Rhizosphere

Considerable progress has been made during the last decade towards understanding and quantifying the input and turnover of plant carbon in the rhizosphere. This was made possible by the development (partially by the authors) and combination of appropriate new methods, such as:

• –homogeneous labelling of whole plants with ¹⁴C
• –distinction between root and microbial respiration
• –separation of soil zones of known distances from the roots
• –determination of microbial soil biomass.

These methods were applied to study the following aspects:

• –release of organic plant carbon into the soil by growing roots
• –utilization of this plant carbon by the microbial biomass in the rhizosphere
• –related influence on the turnover of soil organic matter, and
• –spatial range of such root influence in the soil.

About 19% of the total photosynthetic production of the investigated plants was released into the rhizosphere as organic material. Most of this (15%) was transformed by the rhizosphere microorganisms into CO₂, while only a small fraction (4%) remained in the soil, mainly as microbial cells (2.5%). As a result, microbial rhizosphere biomass increased considerably. Relative to the organic C-input, however, the incorporation of root derived carbon by the microbial biomass was remarkably low (13%). Along with the increase in microbial rhizosphere biomass, the presence of plant roots also enhanced the decomposition of soil organic matter and affected soil aggregate stability. Root carbon and root influences were even detected up to 20 mm away from the roots. This may be partially attributed to the contribution of root derived volatiles. Accordingly, both the actual volume of the rhizosphere and its metabolic significance is greater than what has so far been assumed. Possible interactions involving root, soil and microbial carbon are discussed.     

Distribution of exudates of lupin roots in the rhizosphere under phosphorus deficient conditions

Abstract

The distribution of secretory acid phosphatase and organic acids enhanced by phosphorus deficiency in lupin rhizosphere was investigated using a rhizobox system which separated the rhizosphere soil into 0.5 mm fractions. In the soil fraction closest to the root surface, the lupin exudates displayed an acid phosphatase activity of 0.73 u g^?1 dry soil and citrate concentration of 85.2 μmol g^?1 dry soil, respectively. The increase of the acid phosphatase activity-induced an appreciable depletion of organic P in the rhizosphere, indicating that lupin efficiently utilized the organic P from soil through the enzyme activitye The sterile treatments demonstrated that the acid phosphatase in the rhizosphere was mainly derived from lupin root secretions. The secretory organic acids enhanced considerably the solubility of the inorganic P in three types of soil and a sludge. However, the secretory acid phosphatase and organic acids from lupin roots were only detected in a considerable amount in 0-2.5 mm soil fractions from root surface.     

Biodegradation of low molecular weight organic acids in rhizosphere soils from a tropical montane rain forest

Root exudation of organic acids could be an important strategy for plant acquisition of phosphorus (P) from P-deficient soils in tropical rain forests. However, the efficacy of organic acids on P mobilization in the rhizosphere could be reduced due to their rapid biodegradation by rhizosphere microorganisms. To assess the dynamics and function of organic acids in the rhizosphere soils in tropical rain forests, we examined the concentrations of oxalate, citrate, and malate in soil solution and the mineralization kinetics of ¹⁴C-radiolabelled oxalate and citrate in the rhizosphere and bulk soil fractions. We compared two tropical montane rain forests from Mt. Kinabalu, Borneo that share similar parent material (i.e., sedimentary rocks) and climate but differ in terms of soil age. The older soil (Tertiary age materials) was affected by podzolization and had less inorganic labile P compared to the younger soil (Quaternary colluvial deposits). In the P-deficient older soil, the rhizosphere soil solution contained markedly higher concentrations of oxalate, citrate, and malate than did the bulk soil, whereas in the P-rich younger soil, the levels of organic acids in the rhizosphere were lower. The higher levels of organic acids in the rhizosphere of P-deficient soils are caused by greater root exudation and the lower sorption capacity for organic acids. The results of mineralization kinetics showed that oxalate and citrate in soil solution were rapidly mineralized in both rhizosphere and bulk fractions of both P-rich and P-deficient soils, having short mean residence times (2.3–13.1 h for oxalate and 0.8–1.6 h for citrate). The mineralization rates of oxalate and citrate were highest in the rhizosphere fraction of the P-deficient soil, where the pool of organic acids was largest and rapidly replenished by root exudation. Our data indicate that consumption as well as production of organic acids in the rhizosphere could be enhanced in P-deficient soil. The efficacy of organic acids on P mobilization in the rhizosphere in tropical montane rain forests appears to vary depending on the level of soil P availability and the anion sorption capacity, attributable to soil aging with podzolization.     

Flavonoids of white lupin roots participate in phosphorus mobilization from soil

The impact of flavonoids released by phosphorus-deficient white lupin roots on inorganic P and soil microorganisms is largely unknown. We report that flavonoids isolated from white lupin roots mobilized inorganic phosphorus and decreased soil microbial respiration, citrate mineralization, and soil phosphohydrolase activities, but did not reduce the soil ATP content. The results suggest that white lupin's release of flavonoids into the rhizosphere plays a significant role in its efficient P-acquisition strategy by solubilizing Fe-bound P and by limiting the microbial mineralization of citrate.     

Effect of plant roots on carbon metabolism of soil microbial biomass

The utilization of plant- and soil-C by the microbial biomass in the rhizosphere of maize plants was investigated as a function of root proximity. The plants were cultivated in pots with divided root chambers and their shoots supplied with ¹⁴CO₂ for 23 days. Subsequently the individual soil zones were analyzed for organic C, ¹⁴C, biomass C and biomass ¹⁴C. Plant roots induced a 197% increase in microbial biomass and a 5.4% decrease in soil organic C compared with an 1.2% decrease in the unplanted control soil. The contributions of plant- and soil-C to this increased microbial growth amounted to 68% and 32% respectively. Biomass-¹⁴C corresponded to 1.6% of the total photosynthetically fixed ¹⁴C, to about 15% of the organic ¹⁴C-input into the rhizosphere and to 58% of the plant carbon remaining in soil after the removal of roots. 20% of this biomass-¹⁴C was found outside the immediate root zone. These results demonstrate that growing roots are a significant C-source for the microbial biomass and render an additional fraction of soil-C available to microbial utilization. The efficiency of C-utilization by the rhizosphere biomass is lower than values obtained with liquid cultures in laboratory experiments. The supply of plant-C to the microbial biomass outside the immediate root vicinity indicates that the overall volume of the maize rhizosphere is greater than what has been supposed so far.     

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.     

长期定位施肥对玉米根际土壤团聚体有机碳与钾素含量及其相互关系的影响

  目的  探讨长期定位施肥对玉米根际土壤中不同形态钾素含量变化及其与团聚体组分有机碳含量间的耦合关系,为红壤合理施肥提供理论依据。  方法  依托红壤长期施肥定位试验（始于1986年）,采集不施肥（CK）、氮磷肥（NP）、氮磷钾肥（NPK）、2倍氮磷钾肥（DNPK）和氮磷钾配施有机肥（NPKM）5个处理玉米根际土壤,测定不同粒级土壤团聚体的有机碳、交换性钾和非交换性钾含量,探明它们之间的相关关系。  结果  各施肥处理玉米根际土壤不同粒级团聚体组分中 > 2 mm组分均显著低于0.25 ~ 2 mm和 < 0.25 mm组分,但NPKM处理 > 2 mm的组分最高,< 0.25 mm组分则显著低于其他处理;而各团聚体组分间、同一粒级不同处理间全钾含量均无显著差异。与NP处理相比,NPK处理在 > 2 mm、0.25-2mm和 < 0.25mm团聚体的交换性钾含量分别增加了64.83%、31.12%和32.43%,非交换性钾含量分别提高了11.74%、16.86%和14.54%。相关分析表明,> 2 mm团聚体中有机碳与交换性钾、非交换性钾含量均呈现出显著的正相关关系（R2分别为0.72和0.77,P < 0.05）,且 > 2 mm团聚体中有机碳含量增加0.1 g kg?1,交换性钾和非交换性钾含量分别提高5.69 mg kg?1和 2.37 mg kg?1。  结论  在南方红壤地区,长期有机无机肥配施是提高玉米根际土壤大团聚体比例的重要措施,且大团聚体中有机碳含量的增加促进了交换性钾和非交换性钾含量的提高,可为满足作物根系的钾素需求奠定基础。     

Changes in phosphorus fractions in the rhizosphere of some crop species under glasshouse conditions

The aim of the present study was to evaluate the effect of cultivation of some agronomic plant species on inorganic soil‐P fractions in different sampling zones. The experiment was conducted as a completely randomized design as factorial in three replicates. The factors were plant species (10 different species and control) and soil‐sampling zone (soil adhering to root mat, rhizosphere soil, and nonrhizospheric soil). The thin‐slicing technique was modified and used to sample rhizosphere soil. The percentages of P fractions in the planted and control soils were near 64% apatite (apatite‐P), 24% octa‐Ca‐phosphates (OCP‐P), 7% Fe‐phosphates (Fe‐P), 4% di‐Ca‐phosphates (DCP‐P), 1% Al‐phosphates (Al‐P), and 0% occluded‐P (O‐P). All of the studied plant species decreased significantly all of the inorganic P fractions in soil adhering to root mat and in rhizosphere soil compared to those in nonrhizosphere soil. However, these decreases were not equal for each fraction and the percentage of apatite‐P increased in rhizosphere soil of the plant species. The means of total P, soluble P, DCP‐P, OCP‐P, Al‐P, and Fe‐P were lower in soil adhering to root mat compared to those in rhizosphere soil. However, this difference was only significant for OCP‐P. In contrast, the mean of apatite‐P in soil adhering to root mat was significantly higher than that in rhizosphere soil. The changes of apatite‐P may be more governed by microbial activities (especially mycorrhizal symbiosis) which may be higher in rhizosphere soil compared to soil adhering to root mat.     

Effect of arbuscular mycorrhizal fungi on rhizosphere organic acid content and microbial activity of trifoliate orange under different low P conditions

ABSTRACT

Arbuscular mycorrhizal (AM) fungi can improve plant phosphorus (P) uptake; however, information about how AM fungi affect rhizosphere organic acid and microbial activity to alleviate citrus low P stress is limited. Here, a pot experiment was conducted to evaluate the effect of AM fungi (Rhizophagus intraradices, Ri) inoculation on rhizosphere organic acid content, microbial biomass (MB) and enzyme activity of trifoliate orange (Poncirus trifoliata L. Raf.) seedlings grown under three low P conditions. The results showed that mycorrhizal seedlings all recorded higher P concentrations, plant biomass and better root morphology with more lateral and fine roots, but lower root mass ratios, irrespective of P conditions. Mycorrhizal P absorption contribution did not differ significantly among three P conditions. Mycorrhizal seedling rhizosphere soil exhibited lower organic acid content, soil organic P content and ratio of MB-carbon (C)/MB-P, but higher MB and enzyme activity. Additionally, the main organic acids showed a negative relationship with mycorrhizal colonization rate and hyphal length; however, phosphatase and phytase activity had a significantly positive relationship with MB. Therefore, the results suggest that AM fungi inoculation may help citrus to efficiently utilize organic P source by improving microbial activity under low available P conditions.     

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.     

A COMPARISON OF THE SORPTION OF INORGANIC ORTHOPHOSPHATE AND INOSITOL HEXAPHOSPHATE BY SIX ACID SOILS

A comparison has been made of the sorption of inorganic orthophosphate and inositol hexaphosphate by six acidic surface soils from arable land in north-east Scotland. The sorption of inorganic P increased with increasing quantities added and tended towards a maximum, but was never complete. In contrast, the organic P was completely sorbed up to an added quantity which varied with the soil, and above this level the absolute sorption decreased, probably due to the formation of soluble complexes involving iron and aluminium. The sorption sites were apparently the same for the two P forms and, particularly at high levels of addition, the organic P depressed the sorption of inorganic P. Inorganic P did not depress the sorption-of organic P, which was preferentially adsorbed. The results help to explain the extreme stability of inositol hexaphosphate in these soils.     

Phosphorus forms and enzymatic hydrolyzability of organic phosphorus in soils after 30 years of organic and conventional farming

Lower P‐input levels in organic than conventional farming can decrease soil total and available P, which can potentially be resupplied from soil organic P. We studied the effect of 30 y of conventional and organic farming on soil P forms, focussing especially on organic P. Soil samples (0–20 cm) were taken in a field experiment with a nonfertilized control, two organic systems receiving P inputs as animal manure, and two conventional systems receiving only mineral P or mineral P and manure. Soils were analyzed for total, inorganic, organic, and microbial P, by sequential P fractionation and by enzyme additions to alkaline soil extracts. Samples taken prior to starting the experiment were also analyzed. Average annual P balances ranged from –20 to +5 kg ha^–1. For systems with a negative balance, labile and moderately labile inorganic P fractions decreased, while organic and stable inorganic P fractions were hardly affected. Similar quantities and proportions of organic P extracted with NaOH‐EDTA were hydrolyzed in all soils after addition of an acid phosphatase, a nuclease, and a phytase, and enzyme‐stable organic P was also similar among soils. Thus, neither sequential fractionation nor enzyme addition to alkaline soil extracts showed an effect of the type of applied P (manure vs. mineral) on organic P, suggesting that organic P from manure has largely been mineralized. Thus far, we have no indication that the greater microbial activity of the organic systems resulted in a use of stable P forms.     

Effects of pine roots on microorganisms,fauna, and nitrogen availability in two soil horizons of a coniferous forest spodosol

Summary We investigated the effects of pitch pine seedling roots on extractable N, microbial growth rate, biomass C and N, and nematodes and microarthropods in microcosms with either organic (41% C, 1.14% N) or mineral (0.05% C, 0.01% N) horizon soils of a spondosol. Root quantity was manipulated by varying plant density (0, 1, 2, or 4 seedlings) and rhizosphere soil was separated from non-rhizosphere soil by a 1.2 m mesh fabric. In the rhizosphere of organic soil horizons, moisture, microbial growth rate, biomass C and N, and extractable N declined as root density was increased, but there was little effect on nematodes or microarthropods. High levels of extractable N remained after 5 months, suggesting that N mineralization was stimulated during the incubation. In the rhizosphere of mineral soil horizons, microbial growth rate, and nematode and microarthropod abundances increased at higher root density, and in the absence of roots faunal abundance approached zero. Faunal activity was concentrated in the rhizosphere compared to non-rhizosphere soil. In organic soil horizons, roots may limit microbial activity by reducing soil moisture and/or N availability. However, in mineral soil horizons, where nutrient levels are very low, root inputs can stimulate microbial growth and faunal abundance by providing important substrates for microbial growth. Our results demonstrate a rhizosphere effect for soil fauna in the mineral soil, and thus extends the rhizosphere concept to components of the soil community other than microbes for forest ecosystems. Although our results need to be verified by field manipulations, we suggest that the effects of pine roots on nutrient cycling processes in coniferous forests can vary with soil nutrient content and, therefore, position in the soil profile.     

Influence of the rhizosphere on microbial biomass and recently formed organic matter

We have aimed to quantify the effect of roots on the size of the soil microbial biomass, and their influence on the turnover of soil organic matter and on the extent of the rhizosphere. We sampled a sandy clay loam topsoil from two subplots with different treatment histories. One had a normal arable fertilization record, the other had received only inorganic nitrogen fertilizer but no phosphorus and potassium for 30 years. Glucose labelled with ¹⁴C was added to both samples which were then incubated for 4 weeks before the soil was packed in cylinders and planted with ryegrass. In both soils, microbial biomass at the root surface doubled during the first 8 days. At day 15, the microbial biomass had further increased in the fertile soil, and the rhizosphere effect had extended 2.5 mm into the fertile soil, but to only 1 mm in the infertile soil. The microbial ¹⁴C increased threefold near the roots in the fertile soil as a result of assimilation of previously formed microbial residues, but in the infertile soil there was no increase. There was a close relation between the increase in microbial ¹⁴C and a decrease in ¹⁴C soluble in 2 m KCl, indicating that the microbial residues were more weakly adsorbed in the fertile soil. We conclude that the increased microbial population living near the root surfaces re‐assimilated part of the ¹⁴C‐labelled microbial residues in the fertile soil. In the infertile soil, microbial residues resisted decomposition because they were more strongly sorbed on to soil surfaces.     

Relationships between microbial indices in roots and silt loam soils forming a gradient in soil organic matter

Perennial rye grass (Lolium perenne) was grown in a greenhouse pot experiment on seven soils to answer the question whether the microbial colonisation of roots is related to existing differences in soil microbial indices. The soils were similar in texture, but differed considerably in soil organic matter, microbial biomass, and microbial community structure. Ergosterol and fungal glucosamine were significantly interrelated in the root material. This ergosterol was also significantly correlated with the average ergosterol content of bulk and rhizosphere soil. In addition, the sum of fungal C and bacterial C in the root material revealed a significant linear relationship with microbial biomass C in soil. The colonisation of roots with microorganisms increased apparently with an increase in soil microbial biomass. In the root material, microbial tissue consisted of 77% fungi and 23% bacteria. In soil, the fungal dominance was slightly, but significantly lower, with 70% fungi and 30% bacteria. Fungal glucosamine in the root material was significantly correlated with that in soil (r=0.65). This indicates a close relationship between the composition of dead microbial remains in soil and the living fraction in soil and root material for unknown reasons.     

Mycorrhizal community structure,microbial biomass P and phosphatase activities under Salix polaris as influenced by nutrient availability

Low supply of the nutrients nitrogen (N) and phosphorus (P) limit plant growth and spreading, and increase the plant-microbial nutrient competition in subarctic and arctic regions. We investigated the mycorrhizal community structure of a polar shrub willow (Salix polaris) and the microbial turnover in its rhizosphere to explore the adaptation of a mycorrhizal plant in the subarctic tundra. The ectomycorrhizal colonisation ranged from 35 to 64% of the fine root tips and decreased with an increasing soil C/N ratio. In total, 16 ectomycorrhizal morphotypes were found under S. polaris (eight to 13 morphotypes per site, five morphotypes at all four sites). Cenococcum sp. was the most common EM fungus (32% of the ectomycorrhizal fine roots). The abundance of Cenococcum sp. increased with an increasing organic matter content and N/P ratio in the soil. Arbuscular mycorrhizal colonisation of S. polaris was absent or less than 1% of the fine root length. Microbial biomass P accounted for 21–75% of the organic soil P and 6–49% of the total soil P. Microbial biomass P, alkaline and acid phosphatase activities in the rhizosphere increased with increasing soil N concentration. We conclude that a higher N supply decreases the diversity in the mycorrhizal community on polar willows and increases the role of P turnover from the soil microbial biomass for the nutrient supply.     

Distribution of phosphatase activity and various bacterial phyla in the rhizosphere of Hordeum vulgare L. depending on P availability

Despite the importance of the rhizosphere for nutrient turnover, little is known about the spatial patterns of organic phosphorus mineralization by plants and by microorganisms in the rhizosphere. Therefore, the distribution of acid and alkaline phosphatase activity and the abundance of bacteria belonging to various bacterial phyla were investigated in the rhizosphere of barley (Hordeum vulgare L.) as dependent on the availability of inorganic P. For this purpose, we conducted a greenhouse experiment with barley growing in inclined boxes that can be opened to the bottom side (rhizoboxes), and applied soil zymography and fluorescence-in situ-hybridization (FISH). Acid phosphatase activity was strongly associated with the root and was highest at the root tips. Due to P fertilization, acid phosphatase activity decreased in the bulk soil, and less strongly in the rhizosphere. Alkaline phosphatase activity, i.e., microbial phosphatase activity was high throughout the soil in the control treatment and was reduced due to inorganic P fertilization especially in the rhizosphere and less strongly in the bulk soil. P-fertilization slightly increased the total number of bacteria in the rhizosphere. Moreover, P-fertilization decreased the abundance of Firmicutes and increased the abundances of Beta- and Gamma-Proteobacteria. The total number of bacterial cells was significantly higher at the root surface than at the root tip and at a distance of 30 μm from the root surface. Our results show that alkaline phosphatase activity decreased more strongly in the rhizosphere than in the bulk soil due to P fertilization, which might be because of greater C deficiency in the bulk soil compared to the rhizosphere. Furthermore, the results indicate a spatial separation between hotspots of acid phosphatase activity and hotspots of bacteria in the rhizosphere of H. vulgare. Taken together, our study shows that bacteria and phosphatase activity were very heterogeneously distributed in soil, and that the effects of P fertilization on phosphatase activity differed strongly between bulk soil and rhizosphere as well as between various zones of the rhizosphere.     

Growth,phosphorus uptake,and rhizosphere microbial‐community composition of a phosphorus‐efficient wheat cultivar in soils differing in pH

In a pot experiment, the P‐efficient wheat (Triticum aestivum L.) cultivar Goldmark was grown in ten soils from South Australia covering a wide range of pH (four acidic, two neutral, and four alkaline soils) with low to moderate P availability. Phosphorus (100 mg P kg^–1) was supplied as FePO₄ to acidic soils, CaHPO₄ to alkaline, and 1:1 mixture of FePO₄ and CaHPO₄ to neutral soils. Phosphorus uptake was correlated with P availability measured by anion‐exchange resin and microbial biomass P in the rhizosphere. Growth and P uptake were best in the neutral soils, lower in the acidic, and poorest in the alkaline soils. The good growth in the neutral soils could be explained by a combination of extensive soil exploitation by the roots and high phosphatase activity in the rhizosphere, indicating microbial facilitation of organic‐P mineralization. The plant effect (soil exploitation by roots) appeared to dominate in the acidic soils. Alkaline phosphatase and diesterase activities in acidic soils were lower than in neutral soils, but strongly increased in the rhizosphere compared with the bulk soil, suggesting that microorganisms contribute to P uptake in these acidic soils. Shoot and root growth and P uptake per unit root length were lowest in the alkaline soils. Despite high alkaline phosphatase and diesterase activities in the alkaline soils, microbial biomass P was low, suggesting that the enzymes could not mineralize sufficient organic P to meet the demands of plants and microorganisms. Microbial‐community composition, assessed by fatty acid methylester (FAME) analysis, was strongly dependent on soil pH, whereas other soil properties (organic‐C or CaCO₃ content) were less important or not important at all (soil texture).     

Interaction between rhizosphere microorganisms and plant roots: 13C fluxes in the rhizosphere after pulse labeling

The input dynamics of labeled C into pools of soil organic matter and CO₂ fluxes from soil were studied in a pot experiment with the pulse labeling of oats and corn under a ¹³CO₂ atmosphere, and the contribution of the root and microbial respiration to the emission of CO₂ from the soil was determined from the fluxes of labeled C in the microbial biomass and the evolved carbon dioxide. A considerable amount of ¹³C (up to 96% of the total amount of the label found in the rhizosphere soil) was incorporated into the biomass of the rhizosphere microorganisms. The diurnal fluctuations of the labeled C pools in the microbial biomass, dissolved organic carbon, and CO₂ released in the rhizosphere of oats and corn were related to the day/night changes, i.e., to the on and off periods of the photosynthetic activity of the plants. The average contribution of the corn root respiration (70% of the total CO₂ emission from the soil surface) was higher than that of the oats roots (44%), which was related to the lower incorporation of rhizodeposit carbon into the microbial biomass in the soil under the corn plants than in the soil under the oats plants.     

Effects of poultry manure amendment on phosphorus uptake by ryegrass,soil phosphorus fractions and phosphatase activity

Poultry manure (PM) contains a large proportion of phosphorus (P) in mineral-associated forms that may not be readily available for plant uptake. In addition, PM application influences both chemical and biotic processes, and can affect the lability of native soil P. To investigate the effects of PM on soil P availability, we grew ryegrass (Lolium perenne) in greenhouse pots amended with poultry manure. Biomass was harvested at 4, 8, and 16 weeks following PM application, with soil separated into rhizosphere and bulk fractions. Soil was sequentially extracted by H₂O, 0.5 M NaHCO₃, 0.1 M NaOH, and 1 M HCl, and inorganic P (P_i) and enzymatically hydrolyzable organic P (P_oe) were quantitated. Root P concentrations were 37% higher and total P uptake 59% higher with PM application than Control. At week 16, there was 30% more labile-P_i (H₂O- plus NaHCO₃-P_i) in the rhizosphere with PM than in Control. Phosphodiesterase activity increased with PM application. Furthermore, acid phosphomonoesterase, alkaline phosphomonoesterase, and phosphodiesterase activities were all higher in the rhizosphere than in bulk soil at week 16 with PM, indicating that increased labile-P_i was due primarily to stimulation of soil phosphatases to mineralize NaOH-P_oe. Soil pH increased with PM application and plant growth, and may have promoted P availability by decreasing sorption of Al- and Fe-associated inorganic and organic phosphates. These results demonstrate that whereas PM application may initially increase NaOH and HCl-P_i, these fractions can be readily changed into labile-P and do not necessarily accumulate as stable or recalcitrant P in soil.