不同耐盐植物根际土壤盐分的动态变化

以甘肃秦王川引大灌区盐渍化土壤为研究背景,用盆栽根袋法对4种耐盐植物根际和非根际土壤pH和盐分离子的动态变化进行了分析比较。结果表明:4种待测植物随着培养时间的延长土壤pH和EC值呈降低趋势。新疆大叶(Medicago Sativa L.cv.Xinjiangdaye)、向日葵(Helianthus annuus)和霸王(Zygophyllum xanthoxylum)生长90 d后根际土壤pH明显低于非根际,而裸麦(Hordeum vulgare var. vulgare)根际较非根际pH差异不大。霸王和新疆大叶根际土壤EC值较非根际高,而裸麦和向日葵的根际与非根际差异不大。4种供试植物根际K⁺均出现亏缺,Ca²⁺、Na⁺、Mg²⁺、SO^2-₄和Cl^-在新疆大叶、霸王和向日葵3种植物根际均出现富集,对于裸麦:Ca²⁺、Mg²⁺和SO^2-₄ 3种离子在植物根际富集,而Cl^-和Na⁺在根际亏缺。随着待测植物培养时间的增加Na⁺/K⁺、Na⁺/Ca²⁺和Na⁺/Mg²⁺ 这3个比值呈降低趋势,说明Na⁺相对于K⁺、Ca²⁺和Mg²⁺的含量降低,生物措施对Na⁺的移除效果较显著。     

Influence of mature Douglas fir roots on the solid soil phase of the rhizosphere and its solution chemistry

Plants can induce significant changes in the rhizosphere through the uptake of water and ions, the exudation of organic compounds and the activities of micro-organisms. The aim of the present study was to assess the influence of tree roots on the chemistry (pH, exchangeable cations, total organic carbon) of both the solid phase of the soil and the soil solutions, extracted by centrifugation, under a mature Douglas fir stand over two distinct seasons (March and September, 1999). The chemical characteristics of either the solid soil phase or the soil solutions of the rhizosphere were found to be different from those of the bulk soil. The cation exchange capacity, base saturation and organic C were all greater in the rhizosphere than in the bulk soil, as a possible result of rhizodeposition, incorporation of decaying root material and micro-organism activity. The concentration of all elements increased in the rhizosphere solutions as compared to the bulk soil solutions, except for P. The pH was lower in the rhizosphere than in the bulk soil for both the solid soil phase and the soil solutions. Despite the greater overall Al concentration of the rhizosphere solutions, as compared to the bulk soil solutions, we suggest that in both, Al toxicity was efficiently restricted by both high Ca + Mg contents and Al complexation with various ligands.     

Origins of root-mediated pH changes in the rhizosphere and their responses to environmental constraints: A review

The aim of the present review is to define the various origins of root-mediated changes of pH in the rhizosphere, i.e., the volume of soil around roots that is influenced by root activities. Root-mediated pH changes are of major relevance in an ecological perspective as soil pH is a critical parameter that influences the bioavailability of many nutrients and toxic elements and the physiology of the roots and rhizosphere microorganisms. A major process that contributes root-induced pH changes in the rhizosphere is the release of charges carried by H⁺ or OH^– to compensate for an unbalanced cation–anion uptake at the soil–root interface. In addition to the ions taken up by the plant, all the ions crossing the plasma membrane of root cells (e.g., organic anions exuded by plant roots) should be taken into account, since they all need to be balanced by an exchange of charges, i.e., by a release of either H⁺ or OH^–. Although poorly documented, root exudation and respiration can contribute some proportion of rhizosphere pH decrease as a result of a build-up of the CO₂ concentration. This will form carbonic acid in the rhizosphere that may dissociate in neutral to alkaline soils, and result in some pH decrease. Ultimately, plant roots and associated microorganisms can also alter rhizosphere pH via redox-coupled reactions. These various processes involved in root-mediated pH changes in the rhizosphere also depend on environmental constraints, especially nutritional constraints to which plants can respond. This is briefly addressed, with a special emphasis on the response of plant roots to deficiencies of P and Fe and to Al toxicity. Finally, soil pH itself and pH buffering capacity also have a dramatic influence on root-mediated pH changes.     

Effects of ammonium sulphate application on the chemistry of bulk soil,rhizosphere, fine roots and fine-root distribution in a Picea abies (L.) karst. stand

The effect of ammonium sulphate application on the bulk and rhizosphere soil chemistry, elemental concentration of living fine roots (<2 mm in diameter), amounts of living and dead fine roots, root length density and specific root length density were investigated in a 28 year old Norway spruce stand in SW Sweden. The treatments started in 1988. Core samples of the LFH layer and mineral soil layers were sampled in control (C) and ammonium sulphate (NS) treatment plots in 1988, 1989 and 1990. Soil pH and NO₃-S and SO₄-S, Al, Ca, Mg, Mn and K concentrations were measured for both the bulk soil and rhizosphere soil.The pH-values of the bulk and rhizosphere soil decreased in 1989 and 1990 in NS plots compared to control plots, while the SO₄-S concentration increased. The Ca, Mg and K concentration increased in the NS treatment in almost all layers in the bulk and the rhizosphere soil. Ammonium ions may have replaced these elements in the soil organic matter. The NS treatment reduced Mg concentration in fine roots in all layers in 1990. The Al concentrations in the rhizosphere and bulk soil were higher in NS plots in all layers, except at 0–10 cm depth, both in 1989 and 1990. The Al content of living fine roots was higher in NS plots than C plots but the differences were not significant. The NS addition did not affect the P and K contents of fine roots in any soil layer, but the S concentrations of fine roots were significantly higher in NS plots in 1989 and 1990. The fine root necromass was higher in NS than in C in 1990, in the LFH layer, indicating a gradual decrease in the vitality of the fine roots. It was suggested that the NS treatment resulted in displacement of Mg and K from exchange sites in the LFH layer leading to leaching of these cations to the mineral soil. Further application of ammonium sulphate may damage the fine roots and consequently adversely affect the water and nutrient uptake of root systems.     

Microscale fumigation-extraction and substrate-induced respiration methods for measuring microbial biomass in barley rhizosphere

Changes in microbial biomass in the rhizosphere of young barley seedlings was studied. A fumigation-extraction (FE) method with measurement of ninhydrin-reactive nitrogen (NR-N) and a substrate-induced respiration (SIR) method were applied on a microscale to rhizosphere soil samples of approximately 0.1 g. Rhizosphere soil was defined as the soil adhering to the roots when they were carefully separated from the bulk soil. The rhizosphere soil was gently washed off the roots with either distilled water (FE) or with glucose solution (SIR). Shaking and mild sonication was used to disperse the soil without disrupting the roots. Fumigation was carried out by direct addition of liquid chloroform to the isolated soil. These techniques were proven to give reliable results under the experimental conditions of this investigation. Rhizosphere soil was isolated from segments of the roots representing different distances to the seed different root ages. In the rhizosphere of young barley seedlings, biomass NR-N increased significantly compared to the bulk soil from day 6 after sowing (average increases of 33–97%), especially where adventitious roots had developed. From this time, SIR rates were also significantly higher in the rhizosphere than in bulk soil (average increases 72–170%). The average ratio of SIR rate to biomass NR-N was found to be approximately 50% higher in the rhizosphere than in the bulk soil, which may indicate that a larger fraction of the microbial community is potentially active in the rhizosphere as compared to the bulk soil.     

Rhizosphere effect and fine-root morphological adaptations in a chronosequence of silver birch stands on reclaimed oil shale post-mining areas

Mining activities create wastelands that require reclamation. The relief of abandoned opencast oil shale mining area is rugged, and the mining spoil is extremely stony and alkaline (pH 8), with low N and organic content. Planting of fast-growing deciduous tree species such as silver birch (Betula pendula) on post-mining area is the best means to accelerate the development of a new forest ecosystem in such harsh conditions. A chronosequence of silver birch stands (1, 2, 3, 5, 29, 40 years old) was investigated to reveal changes in bulk soil (S) and rhizosphere (R) properties, in rhizosphere effect on bacterial activity and diversity, and in fine-root morphological adaptations in relation to stand development. The rhizosphere effect on bacterial activity was measured as a rhizosphere/soil (R/S) ratio and on species diversity as a similarity (%) between rhizosphere and bulk soil bacterial communities. Bacterial species diversity was determined by denaturing gradient gel electrophoresis (DGGE) technique and was expressed as Shannon diversity index. Biolog EcoPlates were used to determine the summed activity of cultivable bacteria in rhizosphere and bulk soil. Short-root morphological parameters were measured using WinRHIZO™ Pro.Soil pH and available P concentration decreased logarithmically, and N% and organic matter concentration increased linearly with increasing stand age. During the first 30 years of stand development SIR increased an order, from 0.18 to 1.90 mg C g⁻¹. Bulk soil bacterial diversity increased logarithmically with stand age. The bacterial diversity was higher in rhizosphere than in bulk soil. Rhizosphere effect on bacterial activity was low a year after planting, increased more than two times in the next 2 years, and decreased thereafter rapidly with stand age. Rhizosphere effect, indicating plant support to rhizosphere microbial communities, was highest when soil conditions were still poor, but trees had already overcome the transplant shock. All short-root morphological parameters showed certain trends with age. Specific short-root length varied between 56 and 313 m g⁻¹ and decreased logarithmically with stand age and soil improvement. The fastest changes in short-root morphology, rhizosphere effect, and soil pH occurred during the early development of silver birch stands - in the first 5 years; P nutrition and N use efficiency improved simultaneously. Rhizosphere effect and short-root morphological adaptation have an important role in soil and stand development on oil shale post-mining area, and silver birch is a promising tree species for reclamation of alkaline mining spoil.     

Relative contributions of soil chemistry, plant physiology and rhizosphere induced changes in speciation on Ni accumulation in plant shoots

The aim of this study is to rank the relative importance of soil properties, root uptake and root-to-shoot redistribution on the transfer of the trace element nickel from soil to the shoots of non hyperaccumulatings plants. Two contrasting soils and seven plant species have been studied using the radioactive isotope, ⁶³Ni. Shoots and roots were analysed separately and the specific activity of each plant has been measured. The isotopic exchange properties of rhizosphere soil where compared with control non rhizosphere soil. Possible changes in Ni speciation in the rhizosphere have been assessed by comparing the isotopic exchange properties of the rhizosphere and control soil and by comparing the specific activities of Ni in each plant. The capacity of soil to immobilise added radiotracer largely determines root uptake, leading to between a 4- and 40-fold difference between soils for a given species. The redistribution of nickel from roots to shoots was fairly constant for plants grown on the rendzina, but varied strongly between species for the acid soil. This variation enhanced the contrast between species of the soil-to-shoot transfer factor. Root action significantly enhanced immobilisation of added nickel in an acid soil due to the modification of speciation of initially non exchangeable soil nickel, but had little effect on a neutral rendzina. Changes in rhizosphere pH were similar on the two soils. In the acid soil, these pH changes were accompanied by changes in Ni speciation but a causative link has not been established. In the neutral soil pH changes may have modified root uptake properties.     

Dynamics of soil water content in the rhizosphere

Water flow from soil to plants depends on the properties of the soil next to roots, the rhizosphere. Although several studies showed that the rhizosphere has different properties than the bulk soil, effects of the rhizosphere on root water uptake are commonly neglected. To investigate the rhizosphere’s properties we used neutron radiography to image water content distributions in soil samples planted with lupins during drying and subsequent rewetting. During drying, the water content in the rhizosphere was 0.05 larger than in the bulk soil. Immediately after rewetting, the picture reversed and the rhizosphere remained markedly dry. During the following days the water content of the rhizosphere increased and after 60 h it exceeded that of the bulk soil. The rhizosphere’s thickness was approximately 1.5 mm. Based on the observed dynamics, we derived the distinct, hysteretic and time-dependent water retention curve of the rhizosphere. Our hypothesis is that the rhizosphere’s water retention curve was determined by mucilage exuded by roots. The rhizosphere properties reduce water depletion around roots and weaken the drop of water potential towards roots, therefore favoring water uptake under dry conditions, as demonstrated by means of analytical calculation of water flow to a single root.     

Distribution of cutin and suberin biomarkers under forest trees with different root systems

Background and aims

The rhizosphere, the soil immediately surrounding roots, provides a critical bridge for water and nutrient uptake. The rhizosphere is influenced by various forms of root–soil interactions of which mechanical deformation due to root growth and its effects on the hydraulics of the rhizosphere are the least studied. In this work, we focus on developing new experimental and numerical tools to assess these changes.

Methods

This study combines X-ray micro-tomography (XMT) with coupled numerical simulation of fluid and soil deformation in the rhizosphere. The study provides a new set of tools to mechanistically investigate root-induced rhizosphere compaction and its effect on root water uptake. The numerical simulator was tested on highly deformable soil to document its ability to handle a large degree of strain.

Results

Our experimental results indicate that measured rhizosphere compaction by roots via localized soil compaction increased the simulated water flow to the roots by 27 % as compared to an uncompacted fine-textured soil of low bulk density characteristic of seed beds or forest topsoils. This increased water flow primarily occurred due to local deformation of the soil aggregates as seen in the XMT images, which increased hydraulic conductivity of the soil. Further simulated root growth and deformation beyond that observed in the XMT images led to water uptake enhancement of ~50 % beyond that due to root diameter increase alone and demonstrated the positive benefits of root compaction in low density soils.

Conclusions

The development of numerical models to quantify the coupling of root driven compaction and fluid flow provides new tools to improve the understanding of plant water uptake, nutrient availability and agricultural efficiency. This study demonstrated that plants, particularly during early growth in highly deformable low density soils, are involved in active mechanical management of their surroundings. These modeling approaches may now be used to quantify compaction and root growth impacts in a wide range of soils.     

Plant induced alteration in the rhizosphere and the utilisation of soil heterogeneity

Plant-soil interactions result in a special rhizosphere soil chemistry, differing from that of the bulk soil found only a few mm from the root. The aim of this study was to investigate adaptation mechanisms of herbs growing in acid soils through studying their rhizosphere chemistry in a greenhouse experiment and in a field study. Ten herbs were grown in acid soil (pH 4.2 in the soil solution) in the greenhouse. The concentrations of NO₃ ^-, SO₄ ^2-, phosphates, Ca²⁺, Mg²⁺, Mn²⁺, K⁺, Na⁺, NH₄ ⁺ and pH were analysed in soil solutions obtained by centrifugation. The general pattern found was a depletion of nutrients in the rhizosphere compared with their concentrations in the bulk soil. The pH increase (up to 0.7 units) in the rhizosphere soil appeared to be caused by plant uptake of NO₃ ^- (r²=0.88). The ion concentrations in the soil solution of the rhizosphere were dependent on plant species and biomass increase. Although species with a larger biomass and higher growth rates showed a higher degree of ion depletion (except for Na⁺, SO₄ ^2-) in the rhizosphere, there were also species specific responses. A field study of five herbs at five oak forest sites in Southern Sweden (Scania) was also carried out. In addition to the soil solution concentrations, the loss on ignition (LOI) and the concentrations of 0.1 M BaCl₂ extractable K⁺, Mg²⁺, Mn²⁺, Ca²⁺, and Al ions were measured. The amount of soil solution Al was determined as free ionic (quickly reacting) Al. For all species and sites, the LOI and the concentrations of exchangeable cations were higher in the rhizosphere than in the bulk soil, apparently due to the roots preferably growing at organic-rich microsites. The concentrations of the ions as measured in the centrifuged soil solution, were either higher in the rhizosphere than in the bulk soil or were the same in both, except for NO₃ ^- and quickly reacting Al. The lower concentrations of quickly reacting Al in the rhizosphere, compared with the bulk soil could indicate the uptake of Al by the plant or the exudation of complexing substances. The pH differences were only small and mostly non-significant. Plant-soil interactions and the ability of plants to utilise heterogeneity of the soil appear to be more important for plant growth in acid soils than recognised heretofore. Rhizosphere studies provide an important means of understanding plant strategies in acid soils. This revised version was published online in June 2006 with corrections to the Cover Date.     

Water-soluble carbon in roots of rape and barley: impacts on labile soil organic carbon, arylsulphatase activity and sulphur mineralization

Investigating the impact of plant species on sulphur (S) availability in the rhizosphere soil is agronomically important to optimize S fertilization. Bulk, rhizosphere soils and the roots of field-grown rape and barley were sampled 7 times (every fortnight), from March to June, at plant maturity. Root carbon (C) and nitrogen (N) in water extract, along with soil SO₄²⁻-S, labile soil organic-C (HWC) and -N (HWN) in hot water extract, as well as soil arylsulphatase activity were then monitored. The average concentrations of both HWC and HWN were observed in the following decreasing order: rape rhizosphere soil >barley rhizosphere soil >bulk soil. In parallel, the average contents of water extractable-C and -N in rape roots were higher than those in barley roots. These results suggest that soil C and N contents in hot water extract (including rhizodeposition) were correlated with C and N released by roots. Great ARS activities found in rape rhizosphere soil were accompanied by great SO₄²⁻-S mineralization over time. Finally, bulk and rhizosphere soils of rape and barley were pooled from the seven samplings and incubated with the corresponding pooled root water-soluble C of both plant species and glucose-C. After 1 and 9 weeks, a greater net S mineralization (gross mineralization - immobilization) was observed with rape root water-soluble C than with barley root water-soluble C and glucose-C. Conjointly, we found a higher average value of ARS activity in rape rhizosphere than in barley rhizosphere soil. Our findings suggest that plant species, via their rhizodeposition, determine the dynamic of S in soil.     

Changes in water extractable metals,pH and organic carbon concentrations at the soil-root interface of forested soils

Soluble metals are of nutritional and ecotoxicological interest as they are the most readily available form to the biota. Metal solubility in soils is mostly controlled by pH and the organic matter content. The rhizosphere is generally considered as an environment enriched in organic matter and often more acidic (depending on nutritional status of the plant) than the bulk soil. Yet, there is a lack of consensus on the distribution of metals at the soil-root interface. Consequently, the specific objectives of this paper are to compare the chemical properties and the water extractable metal concentrations of the rhizosphere and the bulk soil of forest soil (1) along a gradient in soil contamination and (2) under different tree species. Two study areas were used: (1) Rouyn-Noranda (Canada) where samples were collected along a gradient in metal contamination at a distance of 0.5, 2 and 8 km downwind from a copper smelter; (2) Saint-Hippolyte (Canada) where the effect of three tree species (Abies balsamea, Acer saccharum and Betula papyrifera) was studied. In the field, the rhizosphere was operationally defined as the soil adhering to the roots after agitation, soil falling from the roots and the rest of the soil composing the bulk soil. Once in laboratory, a second agitation was performed to separate the rhizosphere into an inner and an outer component. Water extractable metal concentrations (Al, Ca, Cd, Co, Cr, Cu, Fe, Li, Mg, Mn, Ni, Pb and Zn) were quantified either with an ICP-AES or a GFAAS. Measurements of pH, electrical conductivity (EC), water-extractable organic carbon (WEOC) and solid phase organic carbon (SPOC) were performed. Results systematically indicate that EC, WEOC and SPOC follow the sequence inner rhizosphere > outer rhizosphere > bulk soil. The pH is always lower in the inner rhizosphere than in the bulk soil, while the outer rhizosphere frequently shows an inconstant behaviour. The results also show a clear gradient following inner rhizosphere > outer rhizosphere > bulk soil for water extractable Al, Ca, Cd, Cu, Fe, Mg, Mn, Ni, Pb and Zn. Li, Co and Cr levels were below method detection limit in all cases. WEOC seems to be the main variable related to the water-extractable metals concentrations. The gradient in metal contamination at Rouyn-Noranda was not as expected in the water extracts with the site at 2 km frequently presenting higher metal concentrations than the sites at 0.5 and 8 km. Moreover, a tree species effect did not clearly immerge for any of the chemical properties studied. However, the water extractable Ca concentrations were higher in the soil under Acer saccharum. The effects of the metal gradient and of the tree species may be more pronounced if stronger extractants are used. The addition of an outer rhizosphere component is useful as its behaviour is not consistently intermediate between the inner rhizosphere and bulk soil.     

Changes in the extractability of cations (Ca,Mg and K) in the rhizosphere soil of Norway spruce (Picea abies) roots

Nutrient concentrations in the rhizosphere soil can be higher, lower or remain unchanged compared to the bulk soil, but relatively little is known about such changes for basic cations in the rhizosphere of tree roots. A modified root container technique of studying rhizosphere processes was employed. Plexiglas cylinders were horizontally split by a membrane with 30 M mesh size into an upper compartment for root growth and a root-free lower compartment, each with an inner diameter of 5 cm and a height of 10 cm. One 2-year-old Norway spruce (Picea abies) seedling was transplanted from a nursery into each cylinder. Plants were not specifically inoculated, but roots were colonised by a mix of ectomycorrhizal fungi originating from the nursery. The nutrient poor mineral soil used in the experiment was taken from a forest site in Bayerischer Wald, southern Germany. The soil was either supplied with a mix of Ca, Mg and K, or not supplied with these cations. Plants were harvested 30 weeks after transplanting. The nylon membrane between the root compartments restricted root growth to the upper compartment, so that by the end of the experiment a root mat was formed at the top side of the membrane. In the lower compartment, soil nearest to the root mat was regarded as rhizosphere soil while soil in a distance from the root mat was regarded as bulk soil. In the upper compartment, rhizosphere soil was obtained at the end of the experiment by gently shaking the roots. The soils were analysed for Ca, Mg and K contents following two different soil extraction methods. In the fertilised treatment, H₂O-extractable Ca and Mg were accumulated in the rhizosphere. In contrast, K (NH₄Cl-extraction) was depleted in the rhizosphere. In the bottom tube, the depletion of K (NH₄Cl-extraction) was restricted to 1 cm distance from the root mat. In unfertilised soil, Ca, Mg and K concentrations did not differ clearly between rhizosphere and bulk soils. The results indicated that the occurrence of cation gradients in the rhizosphere depended on the level of soil nutrient supply. Distinct rhizosphere effects were measured by conventional soil extraction methods only when the soil was freshly fertilised with mineral elements prior to the experiment. In this case, K depletion in the rhizosphere reflected higher K uptake by the fertilised Norway spruce plants. For low-nutrient soils, novel techniques are required to follow subtle changes in the rhizosphere.     

Experimental study of solute transport and extraction by a single root in soil

The fate of ¹⁴C-2,4,6-trinitrotoluene ([U-¹⁴C]TNT) in soil/plant systems was studied using onion (Allium cepa L.) plants with only a single root. It was found that the single roots grew exponentially and that the rate of water uptake of the onion plants increased exponentially, as well. The concentration of [U-¹⁴C] in the roots at first increased and then appeared to reach a steady state, while the [U-¹⁴C] concentration in the leaves was found to increase linearly with time. The [U-¹⁴C] concentration in the rhizosphere increased gradually, while in the bulk soil it decreased slowly. The accumulation of [U-¹⁴C] in the rhizosphere is likely to difference between movement into the rhizosphere (through advective mass flow of soil water by root uptake) and its uptake into the roots. The distribution of ¹⁴C in the soil/plant system was found to be 60–85% in the soil solid phase, 7–11% in the soil liquid phase, <1% in the soil air phase, <1% in the root compartment, and <0.01% in the leaf compartment. The maximum RCF (root concentration factor) value for TNT and its derivates was found to be about 20, and the maximum TSCF (transpiration stream concentration factor) was 0.18. These values can be changed by a variety of factors in soil-plant systems     

Three-dimensional visualization and quantification of water content in the rhizosphere

? Despite the importance of rhizosphere properties for water flow from soil to roots, there is limited quantitative information on the distribution of water in the rhizosphere of plants. ? Here, we used neutron tomography to quantify and visualize the water content in the rhizosphere of the plant species chickpea (Cicer arietinum), white lupin (Lupinus albus), and maize (Zea mays) 12 d after planting. ? We clearly observed increasing soil water contents (θ) towards the root surface for all three plant species, as opposed to the usual assumption of decreasing water content. This was true for tap roots and lateral roots of both upper and lower parts of the root system. Furthermore, water gradients around the lower part of the roots were smaller and extended further into bulk soil compared with the upper part, where the gradients in water content were steeper. ? Incorporating the hydraulic conductivity and water retention parameters of the rhizosphere into our model, we could simulate the gradual changes of θ towards the root surface, in agreement with the observations. The modelling result suggests that roots in their rhizosphere may modify the hydraulic properties of soil in a way that improves uptake under dry conditions.     

A technique for studying rhizosphere processes in tree crops: soil phosphorus depletion around camellia (Camellia japonica L.) roots

Rhizosphere studies on tree crops have been hampered by the lack of a satisfactory method of sampling soils at various distances in the rhizosphere. A modified root study container (RSC) technique developed for annual crops, grasses and legumes was used to study the mechanisms by which camellia plants (Camellia japonica L.) utilise soil P in the glasshouse and field. Plants belonging to the Camellia family (e.g. tea) have the ability to utilise P from relatively unavailable native P sources and for this reason camellia plants were selected for this study.In the glasshouse trial, the RSCs were filled with a Recent soil, treated with P fertilisers; North Carolina phosphate rock (NCPR), diammonium phosphate (DAP), mono calcium phosphate (MCP) and single superphosphate (SSP) at 200 g P g^-1 soil. A planar mat of roots was physically separated by a 24 m polyester mesh and the soil on the other side of this mesh was cut into thin slices parallel to the rhizoplane and analysed for pH, and different forms of P (organic, P_o and inorganic, P_i) to understand P depletion at different distances from camellia roots. In the field trial this technique was modified and used to study the rhizosphere processes in mature camellia trees fertilised with only SSP and NCPR.In both field and glasshouse trials, all P fertilisers increased all the bulk soil P fractions except NaOH-P_o over unfertilised soil with the greatest increases being in the H₂SO₄-P_i fraction in the NCPR treatment and NaOH-P_i in the SSP treatment. Resin-P, NaOH-P_i and H₂SO₄-P_i were significantly lower in the rhizosphere soil compared to the bulk soil whereas NaOH-P_o was higher in the rhizosphere soil than in the bulk soil. Plant and microbial P uptake were thought to be the major causes for the low resin-P rather than P fixation by Fe and Al because the NaOH-P_i fraction which is a measure of Fe-P and Al-P, also decreased in the rhizosphere soil. The rhizo-deposition of NaOH-P_o suggests that labile inorganic P was immobilized by rhizosphere microbes which were believed to have multiplied as a result of carbon exudates from the roots. A marked reduction in pH (about 0.2–0.4 in the glasshouse and 0.2 in the field trial) was observed near the rhizoplane compared to that in the bulk soil in all treatments. The pH near the rhizoplane as well as in the bulk soil was highest for NCPR treated soil. The increase in pH in the NCPR treatment over the control was consistent with the number of protons consumed during the dissolution of NCPR. In both trials, the dissolution of NCPR in the rhizosphere was higher than in the bulk soil due to lower pH and plant uptake of solution P in the rhizosphere. The RSC technique proved to be a viable aid to study the rhizosphere processes in tree crops.     

Changes induced by the roots of Erica arborea L. to create a suitable environment in a soil developed from alkaline and fine-textured marine sediments

Background and aims

We report on the modifications induced by the roots of Erica arborea L. on a soil derived from alkaline and fine-textured marine sediments.

Methods

Physical, chemical, mineralogical and biochemical properties of bulk soil and of the rhizosphere of Erica were characterised to evaluate its role on soil development.

Results

Once the upper horizons had been decarbonated because of geomorphic and pedogenic processes, Erica colonised the soil and progressively modified it through the activity of roots. In the upper horizons, there was no difference between rhizosphere and bulk soil for pH, organic C and exchangeable Al and H. At depth, pH, organic C and exchangeable Al and H differed between rhizosphere and bulk soil. The weathering reactions induced by the Erica roots caused a relative quartz enrichment in the rhizosphere compared with the bulk soil. In the E, EB and Bw horizons, the microbial community of the rhizosphere appeared better adapted than in the underlying 2Bw horizons, where the rhizospheric microorganisms were poorly adapted as these horizons represented the boundary between acid and sub-alkaline soil environments.

Conclusions

The activity of Erica roots modified soil properties so to produce more favourable conditions for itself and the rhizosphere microflora.     

Gradients in soil solution composition between bulk soil and rhizosphere – In situ measurement with changing soil water content

Soil solution composition changes with time and distance from the root surface as a result of mass flow, diffusion, plant nutrient uptake and root exudation. A model system was designed, consisting of a root compartment separated from the bulk soil compartment by a nylon net (30 m mesh size), which enabled independent measurements of the change of soil solution composition and soil water content with increasing distance from the root surface (nylon net). K⁺ concentration in the rhizosphere soil solution decreased during the initial growth stage (12 days after planting, DAP). Thereafter K⁺ accumulated with time, due to mass flow as the dominating process. The extend of K⁺ accumulation depended on the initial fertiliser application. As K⁺ concentrations in soil solution increase, not only as a result of transport exceeding uptake, but also as a result of decreasing soil water content, it is hypothesised that K concentration in soil solution is not the only trigger for the activity of K transporters in membranes, but ABA accumulation in roots induced by decreasing soil matric potentials may add to the regulation. A strong decrease of rhizosphere pH with time is observed as a result of H⁺ efflux from the roots in order to maintain cation-anion balance. In addition the K⁺ to Ca²⁺ ratio was altered continuously during the growing period, which has an impact on Ca²⁺ uptake and thus firmness of cell walls, apoplast pH, membrane integrity and activity of membrane transporters. The value of osmotic potential in the rhizosphere soil solution increased with time indicating decreasing soil water availability. Modelling approaches based on the data obtained with the system might help to fill in the time gaps caused by the low temporal resolution of soil solution sampling method.     

Localised nitrate and phosphate application enhances root proliferation by wheat and maximises rhizosphere alkalisation in acid subsoil

The soil pH in the vicinity of the roots can be changed by an imbalance in supply of predominant anions or cations. A soil column experiment examined the effects of localised supply of nitrate and P on plant growth and pH change in a Podosol (pH 3.76 in 0.01 M CaCl₂ and pH buffering capacity 0.81 cmol kg^?1 pH^?1). Nitrate [(Ca(NO₃)₂] and P [(NaH₂PO₄)] fertilizers were applied alone or in combination to either 0–5 cm or 10–15 cm layer of the soil column. Aluminium-tolerant (ET8) and sensitive (ES8) wheat (Triticum aestivum, L) were grown for 38 days. Plant height, water use and tiller number were measured during the growth period. Biomass production, root growth and soil pH were determined at the final harvest. On average, ET8 had a greater shoot biomass, root length and water use than ES8. The greatest shoot biomass and water use were achieved where N and P were applied together in the 0–5 cm layer, followed by N and P together in the 10–15 cm layer and the lowest where N was applied in the 0–5 cm and P in the 10–15 cm layer. Root length density in the subsoil was greatest where N and P were applied together followed by N alone, and the lowest with the supply of P alone. The effect of localised supply was greater on rhizosphere pH than bulk soil pH. The application of N and P together in topsoil and subsoil layers increased rhizosphere pH by 0.4 and 0.5 units respectively, compared to the corresponding layers in the treatment where N and P were applied uniformly in the whole soil column. Changes in rhizosphere pH were similar under both genotypes, although ET8 produced more roots than ES8 in the soil profile. The results suggest that the combined application of nitrate and P is necessary to maximise root proliferation and root-induced alkalisation in acid subsoil.     

Response of microbial activity and biomass in rhizosphere and bulk soils to increasing salinity

Background and aims

The impact of salinity on microbes has been studied extensively but little is known about the response of soil microbial activity and biomass to increasing salinity in rhizosphere compared to bulk (non-rhizosphere) soil.

Methods

Barley was grown for 5 weeks in non-saline loamy sand to which salt (NaCl) was added. The electrical conductivity in the saturated extract (ECe) was 1, 13 and 19 dS m^?1 for non-saline and two saline soils. Pots without plants were prepared in the same manner and placed next to those with plants. The water content in all pots was maintained at 75 % of water-holding capacity by weight. After 5 weeks the planted and unplanted pots were harvested to collect rhizosphere and bulk soil, respectively. The collected soil was then used for an incubation experiment. The EC levels in the pot experiment (EC1, EC13 and EC19, referred to as original) were either maintained or increased by adding NaCl to adjust the EC to 13, 19, 31 and 44 dS m^?1. CO₂ release was measured continuously for 20 days, microbial biomass C (MBC) was measured at the start and the end of the incubation experiment.

Results

In general, cumulative respiration and microbial biomass C concentration in rhizosphere and bulk soil decreased to a similar extent with increasing adjusted EC. However, compared to the treatments where the EC was maintained, the percentage decrease in cumulative respiration when the EC was increased to EC44 was smaller in rhizosphere than in bulk soil.

Conclusion

Overall, the reduction of cumulative respiration with increasing salinity did not differ between rhizophere and bulk soil. But microbes in rhizosphere soil were more tolerant to high EC than those in bulk soil which could be due to the greater substrate availability in the rhizosphere even after the soil was removed from the roots.