Bulk Soil Solution Research Articles

Metal Solubility and Speciation in the Rhizosphere of Lupinus albus Cluster Roots

The objective of this study was to investigate the influence of root exudation of organic acid anions on the speciation of major and trace metal cations in the rhizosphere of Lupinus albus cluster roots. Plants were grown in rhizoboxes containing repacked weakly acidic loam. Bulk soil solutions and, during the lifetime of cluster roots, rhizosphere solutions were collected using micro suction cups. During organic acid anion exudation bursts, metals in the rhizosphere of cluster roots were strongly mobilized. The concentrations of dissolved organic carbon derived from soil organic matter increased parallel to organic acid anions. Speciation calculations revealed that, during exudation, Al, Ca, Mn, and Zn in the cluster root rhizosphere were mainly bound with citrate, while Cu and Pb were always strongly bound to soil-derived dissolved organic matter. Our results indicate that cluster root exudation led on one hand to direct mobilization and complexation of metals like Al, Fe, and Zn by citrate and on the other hand to the mobilization of soil organic matter which complexes and solubilizes Cu and Pb.

The effect of bacteria and fungi on chemical weathering and chemical denudation fluxes in pine growth experiments

Vascular plants and associated microbial communities affect the nutrient resources of terrestrial ecosystems by impacting chemical weathering that transfers elements from primary minerals to other ecosystem pools, and chemical denudation that transports weathered elements out of the system in solution. We performed a year-long replicated flow-through column growth experiment to isolate the effects of vascular plants, ectomycorrhiza-forming fungi and associated bacteria on chemical weathering and chemical denudation. The study focused on Ca2+, K+ and Mg2+, for which the sole sources were biotite and anorthite mixed into silica sand. Concentrations of the cations were measured in input and output solutions, and three times during the year in plant biomass and on exchangeable cation sites of the growth medium. Weathering and denudation fluxes were estimated by mass balance, and mineral surface changes, biofilm and microbial attachments to surfaces were investigated with scanning electron microscopy. Both bacteria and fungi increased weathering fluxes compared to abiotic controls. Without a host plant denudation rates were as large as weathering rates i.e. the weathering to denudation ratio was about one. Based on whole year fluxes, ectomycorrhizal seedlings produced the greatest weathering to denudation ratios (1.5). Non-ectomycorrhizal seedlings also showed a high ratio of 1.3. Both ectomycorrhizal hyphal networks and root hairs of non-ectomycorrhizal trees, embedded in biofilm (microorganisms surrounded by extracellular polymers), transferred nutrients to the host while drainage losses were minimized. These results suggest that biofilms localize both weathering and plant nutrient uptake, isolating the root-hypha-mineral interface from bulk soil solution.

Relação entre potássio na solução do solo, umidade e condutividade elétrica aparente do solo

Através de modelos que relacionam umidade (teta), condutividade elétrica aparente do solo (CEa) e condutividade elétrica da solução do solo (CEw), objetivou-se avaliar, para condição de laboratório, a viabilidade de se estimar a concentração de potássio na solução do solo (K), para solos de classe textural franca (CTf) e franco-arenosa (CTfa). Cinco soluções de cloreto de potássio, referentes a cinco condutividades elétricas (1,0, 2,5, 4,0, 5,5 e 7,0 dS m-1), foram aplicadas sobre o solo acondicionado em colunas de PVC, de forma a se obter cinco teta do solo (17,0 ± 1,4, 19,0 ± 1,8, 21,4 ± 2,2, 23,4 ± 2,2 e 26,0 ± 3,0%, em volume). Efetuaram-se leituras de teta e CEa com um aparelho de reflectometria no domínio do tempo (TDR) e se extraiu solução do solo com extrator de cápsula de cerâmica, para determinar a CEw e o K. Três modelos foram ajustados aos dados, por meio de planilha eletrônica. É viável estimar a concentração de K na solução do solo, a partir detetae CEa, para condição de laboratório, por meio dos modelos de Rhoades et al. (1976), Vogeler et al. (1996) e Mualen & Friedman (1991), adaptados com uma relação entre CEw e K do tipo potência, nas faixas de 0 a 60 e 0 a 120 mg L-1, para solos de CTf e CTfa, respectivamente.

Effects of metals on life cycle parameters of the earthworm Eisenia fetida exposed to field-contaminated, metal-polluted soils

Two control and eight field-contaminated, metal-polluted soils were inoculated with Eisenia fetida (Savigny, 1826). Three, 7, 14, 21, 28 and 42 days after inoculation, earthworm survival, body weight, cocoon production and hatching rate were measured. Seventeen metals were analysed in E. fetida tissue, bulk soil and soil solution. Soil organic carbon content, texture, pH and cation exchange capacity were also measured. Cocoon production and hatching rate were more sensitive to adverse conditions than survival or weight change. Soil properties other than metal concentration impacted toxicity. The most toxic soils were organic-poor (1–10 g C kg −1), sandy soils (c. 74% sand), with intermediate metal concentrations (e.g. 7150–13,100 mg Pb kg −1, 2970–53,400 mg Zn kg −1). Significant relationships between soil properties and the life cycle parameters were determined. The best coefficients of correlation were generally found for texture, pH, Ag, Cd, Mg, Pb, Tl, and Zn both singularly and in multivariate regressions. Studies that use metal-amended artificial soils are not useful to predict toxicity of field multi-contaminated soils.

Selenium Behavior in Open Bulk Precipitation, Soil Solution and Groundwater in Alluvial Fan Area in Tsukui, Central Japan

A monitoring study was carried out in an alluvial fan area in Tsukui, Central Japan during the study period of 1999–2003, in order to explain selenium (Se) behaviors in ecosystem combined with air, soil and groundwater. Monthly Se concentrations in open bulk precipitation (rainfall+aerosol, gaseous deposition and etc.), soil solution (collected by porous ceramic-cup) and groundwater ranged from 0.1 to 1.4 μg L−1 (volume-weighted average: 0.34 μg L−1), 0.21 to 1.0 μg L−1 (0.48 μg L−1) and 1.6 to 2.4 μg L−1 (2.2 μg L−1), respectively. Se concentration in open bulk precipitation was negatively correlated with the rainfall amount. Se concentration in soil solution significantly increased with DOC concentration in soil solution. Besides, despite atmospheric Se input and rainfall to the grassland study area, Se concentration in soil solution and groundwater received no significant effect from the rainfall amount, pH, Se, DOC, SO42−, NO3− and EC in rainfall. Even though Se concentrations in groundwater were significantly correlated with soil solution volume, Se, DOC and NO3− and groundwater level, the result of multiple regression analyses (MRA) indicated that the groundwater Se was negatively influenced by groundwater level, which depended on groundwater recharge. Se was transported into the groundwater through the groundwater recharge that largely increased in this alluvial fan study area after heavy rain.

Modified micro suction cup/rhizobox approach for the in-situ detection of organic acids in rhizosphere soil solution

Root–soil interactions can strongly influence the soil solution chemistry in the rhizosphere. In the present study we propose a modification of the classical rhizobox/micro suction cup system to make it suitable for the collection and analysis of organic acids in the rhizosphere. In order to show the potential of the method, we tested the modified system with Lupinus albus L. as a model plant known to exude large amounts of citrate. The suction cups were installed through the transparent front plate of the rhizoboxes just after the emergence of cluster roots in order to allow optimal localized collection of soil solution. A small dead-volume allowed almost immediate stabilisation with formaldehyde of the sampled soil solutions in the collection container to prevent microbial degradation. The concentrations of organic acids were significantly larger in the rhizosphere soil solution of active cluster roots of Lupinus albus L. than in the bulk soil solution (about 400 µM of citrate versus <0.05 µM). We were able to follow the exudation process in-situ, which occurred during 2–3 days. Also the concentrations of other organic acids and inorganic anions differed between the bulk soil and the rhizosphere of cluster roots, normal roots, and nodules.

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.

Modelling 137Cs uptake in plants from undisturbed soil monoliths

A model predicting 137Cs uptake in plants was applied on data from artificially contaminated lysimeters. The lysimeter data involve three different crops (beans, ryegrass and lettuce) grown on five different soils between 3 and 5 years after contamination and where soil solution composition was monitored. The mechanistic model predicts plant uptake of 137Cs from soil solution composition. Predicted K concentrations in the rhizosphere were up to 50-fold below that in the bulk soil solution whereas corresponding 137Cs concentration gradients were always less pronounced. Predictions of crop 137Cs content based on rhizosphere soil solution compositions were generally closer to observations than those based on bulk soil solution composition. The model explained 17% (beans) to 91% (lettuce) of the variation in 137Cs activity concentrations in the plants. The model failed to predict the 137Cs activity concentration in ryegrass where uptake of the 5-year-old 137Cs from 3 soils was about 40-fold larger than predicted. The model generally underpredicted crop 137Cs concentrations at soil solution K concentration below about 1.0 mM. It is concluded that 137Cs uptake can be predicted from the soil solution composition at adequate K nutrition but that significant uncertainties remain when soil solution K is below 1 mM.

Depletion of macro-nutrients from rhizosphere soil solution by juvenile corn, cottonwood, and switchgrass plants

In situ sampling of rhizosphere solution chemistry is an important step in improving our understanding of soil solution nutrient dynamics. Improved understanding will enhance our ability to model nutrient dynamics and on a broader scale, to develop effective buffers to minimize nutrient movement to surface waters. However, only limited attention has been focused on the spatial heterogeneity and temporal dynamics of rhizosphere solution, and still less is known about how rhizosphere solution chemistry varies among plant species. Nutrients in rhizosphere soil solution and changes in root morphology of juvenile corn (Zea mays L. cv. Stine 2250), cottonwood (Populus deltoids L.), and switchgrass (Panicum virgatum L.) were monitored using mini-rhizotron technology. Plants were grown for 10 days in a fine-silty, mixed, superactive, mesic Cumulic Hapludoll (Kennebec series). Micro-samples (100–200 μL) of rhizosphere and bulk soil solution were collected at 24-h intervals at a tension of −100 kPa and analyzed for P, K, Ca, and Mg concentration using Capillary Electrophoresis techniques. Plants were harvested at the end of the 10-day period, and tissue digests analyzed for nutrient content by Inductively Coupled Plasma Spectroscopy. Corn plants produced roots that were 1.3 times longer than those of cottonwood, and 11.7 times longer than those of switchgrass. Similar trends were observed in number of root tips and root surface area. At the end of 10 days, rhizosphere solution P and K concentrations in the immediate vicinity of the roots (<1 mm) decreased by approximating 24 and 8% for corn, and 15 and 5% for cottonwood. A rhizosphere effect was not found for switchgrass. After correction for initial plant nutrient content, corn shoot P, K, and Mg were respectively 385, 132, and 163% higher than cottonwood and 66, 37, and 10% higher than switchgrass. Cottonwood shoot Ca concentration, however, was 68 to 133% higher than that of corn or switchgrass. There was no difference in root P concentration among the three species. Nutrient accumulation efficiency (μg nutrient mm−1 root length) of cottonwood was 26 to 242% higher for P, 25 to 325% higher for Ca, and 41 to 253% higher for Mg than those of corn and switchgrass. However, K accumulation efficiency of corn was four to five times higher than that of the cottonwood and switchgrass. Nutrient utilization efficiency (mg of dry weight produced per mg nutrient uptake) of P, K, and Mg was higher in cottonwood than in corn and switchgrass. These differences are element-specific and depend on root production and morphology as well as plant nutrient status. From a practical perspective, the results of this study indicate that potentially significant differences in rhizosphere solution chemistry can develop quickly. Results also indicate that cottonwood would be an effective species to slow the loss of nutrients in buffer settings.

In situ dynamics of phosphorus in the rhizosphere solution of five species.

Root activity can modify the chemistry of the rhizosphere and alter phosphorus (P) availability and uptake. However, until recently, relatively little was known about the dynamics of soil solution P at the root surface because of our inability to measure in situ changes in solution P at the plant root. A mini-rhizotron experiment with corn (Zea mays L. cv. Stine 2250), soybean [Glycine max (L.) Merr. cv. Pioneer 3563), cottonwood (Populus deltoids L.), smooth brome (Bromus inermis Leyss.), and switchgrass (Panicum virgatum L.) was conducted to measure the spatial and temporal dynamics of P in the rhizosphere solution of a fine silty, P-rich calcareous soil (solid-phase total P concentration = 62 mg kg(-1), pH = 7.68) from western Iowa. Micro-suction cups were used to collect samples of soil solution from defined segments of the rhizosphere, and capillary electrophoresis (CE) was used to determine the P concentration of the soil solution. At the end of 10 d, a decreasing P concentration gradient in soil solution toward the root was observed in corn, cottonwood, and smooth brome. No clear rhizosphere effect was observed for soybean and switchgrass. Statistical analysis indicated significantly lower solution P concentrations in the rhizospheres of corn (p = 0.05), cottonwood (p = 0.01), and smooth brome (p = 0.01) compared with bulk soil solution. Results indicate that P depletion from rhizosphere soil solution depends on plant species. Under the conditions of this study, corn, cottonwood, and smooth brome were more effective in depleting solution P than soybean and switchgrass.

Redox potential of bulk soil and soil solution concentration of nitrate, manganese, iron, and sulfate in two Gleysols

AbstractWhile the reduction of nitrate‐N, Mn(III,IV), Fe(III), and sulfate‐S in soil has been studied intensively in the laboratory, field research has received only limited attention. This study investigated the relationship between redox potential (EH) measured in bulk soil and concentrations of nitrate, Mn2+, Fe2+, and sulfate in the soil solution of two Gleysols differing in drainage status from the Marsh area of Schleswig‐Holstein, Northern Germany. The soils are silty‐sandy and developed from calcareous marine sediments. Redox potentials were monitored weekly with permanently installed Pt electrodes, and soil solution was obtained biweekly by ceramic suction cups from 10, 30, 60, and 150 cm depth over one year. Median EH at 10, 30, 60, and 150 cm depths was 470, 410, 410, and 20 mV in the drained soil and 500, 480, 30, and –170 mV in the undrained soil, respectively. A decrease in EH below critical values was accompanied in the soil solutions (pH 7.4 to 7.8) by disappearance of nitrate below 0 to 200 mV, appearance of Mn2+ below 350 mV, and Fe2+ below 0 to 50 mV. Both metals disappeared from soil solution after aeration. In the sulfide‐bearing environment of the 150 cm depth of the undrained soil, however, the sulfate concentrations were highest at such EH values at which sulfate should be unstable. This discrepancy was reflected in the fact that at this depth bulk soil EH was about 400 mV lower than soil solution EH (250 mV). When investigating the dynamics of nitrate, Mn, and Fe in soils, bulk soil EH provides semi‐quantitative information in terms of critical EH ranges. However, in sulfidic soil environments the interpretation of EH measured in bulk soil is uncertain.

Aluminum Effects onPicea abiesat Low Solution Concentrations

The usual way to evaluate Al toxicity in forest ecosystems is to consider the concentration of Al 3+ in soil solution. The objective of this paper is to show that Al may affect tree seedlings even under conditions of very low bulk-soil solution concentrations of Al. Picea abies (L.) H. Karst. seedlings, with or without ectomycorrhizal inoculation [Hebeloma crustuliniforme (Bull.: St. Amans) Quel.], were grown in perlite substrate with addition of 0, 0.1, and 0.5 mM Al. The perlite buffered the pH at values > 5 and no soluble Al was detected in the system. However, plant biomass was significantly reduced when 0.5 mM Al was added without fungal inoculation, and Al uptake was 2 to 3 times higher than in the control treatment. Fungal inoculation compensated the growth reduction, but did not reduce Al uptake. These results indicate that spruce seedlings are able to mobilize and take up Al from solid phases.

Applying a solute transfer model to phytoextraction: Zinc acquisition by Thlaspi caerulescens

Phytoextraction is the removal of metals from contaminated soils into harvested plant tissues. The rate of phytoextraction is governed by both soil and plant characteristics. Most effort has focused on identifying appropriate plants for phytoextraction, but the benefits from this effort will be marginal unless the metals are in phytoavailable forms in the rhizosphere. The concentration of a metal in the rhizosphere can be estimated using solute transfer models that incorporate: the metal concentration in the bulk soil solution, the buffer power of the soil, diffusion coefficient for the metal, water movement, root size and morphology, and the rate of entry of metal into the roots. Here a solute transfer model is developed to predict the concentration of Zn in the rhizosphere solution ([Zn]ext) of Thlaspi caerulescens, a hyperaccumulator species that could be exploited for Zn phytoextraction. The model predicts that Zn accumulation by T. caerulescens is sub-optimal when the Zn concentration in the bulk soil solution is <27 μM. Such a high [Zn]ext is rare in contaminated agricultural soils, but is possible in the metalliferous substrates where T. caerulescens is endemic. Sensitivity analyses indicate that Zn diffusion is more important than transpiration-driven mass flow for Zn delivery to the root, implying that management of soil physical and hydrological properties will improve phytoextraction. Sensitivity analyses also imply that strategies to enhance the Zn absorption power of the root will not necessarily be successful for enhancing phytoextraction per se. Thus, research into enhancing Zn availability and mobility in soil will be as important as understanding and manipulating Zn uptake by plants. In general, such models can be used to identify constraints to efficient phytoextraction (whether plant or soil) and to determine whether commercial phytoextraction is feasible.

Aluminum Effects on at Low Solution Concentrations

The usual way to evaluate Al toxicity in forest ecosystems is to consider the concentration of Al3+ in soil solution. The objective of this paper is to show that Al may affect tree seedlings even under conditions of very low bulk-soil solution concentrations of Al. Picea abies (L.) H. Karst. seedlings, with or without ectomycorrhizal inoculation [Hebeloma crustuliniforme (Bull.: St. Amans) Quél.], were grown in perlite substrate with addition of 0, 0.1, and 0.5 mM Al. The perlite buffered the pH at values > 5 and no soluble Al was detected in the system. However, plant biomass was significantly reduced when 0.5 mM Al was added without fungal inoculation, and Al uptake was 2 to 3 times higher than in the control treatment. Fungal inoculation compensated the growth reduction, but did not reduce Al uptake. These results indicate that spruce seedlings are able to mobilize and take up Al from solid phases.

Element budgets of pine stands on lignite and pyrite containing mine soils

To analyze the development of pine ecosystems on lignite and pyrite containing mine soils, four pine stands with ages of 3–35 years were investigated in a chronosequence approach. Bulk precipitation, throughfall and soil solution in depths of 20, 40, 70 and 100 cm were studied over a three-year period to determine element fluxes in these forest ecosystems on extreme acidic and saline soils. Element budgets are controlled by the processes induced by pyrite oxidation such as intensive weathering of primary minerals, precipitation and leaching of secondary phases. Despite low water fluxes, element outputs can reach extraordinary high values due to very high concentrations in the soil solution. Although element outputs decrease drastically with stand age, respectively, site age, they exceed those of comparable pine stands on non-mine sites. Nitrogen release from the lignite fraction represents a special characteristic of the examined substrates. Nitrogen losses beneath the rooted zone can be 34 kg N ha −1 yr −1 in 100 cm depth. Element input and output in the examined ecosystems are far from balance. Closed cycling of nutrients seems to be recognizable in the case of potassium.

Importance of Internal Proton Production for the Proton Budget in Japanese Forested Ecosystems

Annual biogeochemical fluxes (bulk precipitation, throughfall, stem flow, soil solution and vegetation uptake) of inorganic elements were observed in eight cool temperature forested ecosystems in Hokkaido, northern Japan, in order to determine the mechanisms of acid neutralization in Japanese forest ecosystems. We compared our results with the other biogeochemical studies in Japan, north Europe and US from the literature. In many Japanese forests, the internal proton production (IPS) by base cation accumulation into the vegetation was a major proton source, rather than external acidic deposition, and the IPS also affected the base cation fluxes from the mineral. IPS in Japanese forest tended to be larger than that in north Europe and US. Our results suggested that the high acid neutralizing ability of Japanese forests could be attributed to the strong relationship between the base cation buffering of the soil and the larger contribution of IPS as a proton source. acidic deposition|biogeochemical cycling|forest ecosystem|Japan|proton budget

Genotypic effects in phytoavailability of radiocaesium are pronounced at low K intensities in soil

The differences in radiocaesium uptake between species were analysed in a series of solution culture and pot trials. Since radiocaesium uptake is very sensitive to the solution potassium (K) concentration, it was hypothesised that species depleting K in the rhizosphere to a larger extent, will have a higher radiocaesium uptake. Five species (bean, lettuce, winter barley, ryegrass and bentgrass) were grown for 18–21 days in nutrient solution spiked with 137Cs and at 4 K concentrations between 0.025 and 1.0 mM. Shoot 137Cs activities all decreased between 17- and 81-fold with increasing K supply. Shoot 137Cs activities were 4-fold different between species at the lowest K supply and 3.4-fold different at high K supply. The same five species were grown in two 134Cs spiked soils with contrasting exchangeable K but similar clay content. Shoot 134Cs activities were up to 19-fold higher in the soil with lowest exchangeable K. Differences in shoot activity concentrations between the species were only 4.5-fold in the high K soil, but were 15-fold in the low K soil. Bulk soil solution 134Cs and K concentration data were combined with radiocaesium uptake characteristics measured in solution culture to predict radiocaesium uptake from soil. Predictions were within 1.6-fold of observations in the high K soil but largely underestimated 134Cs uptake in lettuce, ryegrass and barley in the low K soil. A solute transport model was used to estimate K and radiocaesium concentrations in the rhizosphere. These calculations confirmed the assumption that higher radiocaesium uptake is found for species that deplete K in the rhizosphere to a larger extent.

A mathematical model of root uptake of cations incorporating root turnover, distribution within the plant, and recycling of absorbed species

SummaryUnderstanding the movement of cations in soil, particularly trace metals, is required in many applications such as phytoremediation and pollution control. A dynamic mechanistic model has been developed to describe the long‐term root uptake of a surface‐applied, strongly adsorbed, pollutant metal cation, such as radiocaesium, from soil. It consists of two submodels. The first calculates uptake per unit root length at a local scale over a root's lifetime, for various initial conditions. The second calculates cumulative uptake at a whole‐plant scale for the entire rooting depth as a function of time. The model takes into account the renewal of roots which are considered to have a limited lifetime. Root density may be a function of soil depth and a proportion of roots need not contribute to uptake. Recycling from decaying, or grazed, roots and shoots is considered.Simulations show that removal of cations from soil is exaggerated unless some recycling by roots or shoots is considered or the entire root length does not contribute to uptake. Because of root turnover, uptake is not rapidly limited by diffusive flux of the cation from the bulk soil solution to the solution–root interface. Uptake is very sensitive to root architecture and plant physiology.

Do plants drive podzolization via rock-eating mycorrhizal fungi?

Weathering and supply of nutrients derived from minerals to plants is known to be stimulated by plant symbiotic mycorrhizal fungi. Nutrients are generally thought to pass the bulk soil solution before plant uptake. Jongmans et al. [Jongmans, A.G., van Breemen, N., Lundström, U.S., van Hees, P.A.W., Finlay, R.D., Srinivasan, M., Unestam, T., Giesler, R., Melkerud, P.-A., Olsson, M., 1997. Rock-eating fungi. Nature, 389, 682–683] showed that (ectomycorrhizal) fungi drill innumerable narrow cylindrical pores (diameter 3–10 μm) into weatherable minerals in podzol E horizons. The fungi probably form micropores by exuding strongly complexing low-molecular weight organic acids at their hyphal tips, causing highly local dissolution of Al silicates. Micropores occurred in all thin sections of podzols under Pinus sylvestris and Picea abies available from Sweden (3), Finland (2), Switzerland (2), Denmark (2) and the Netherlands (3), but not in the few available thin sections of non-podzolic soils under broadleaves. Many weatherable minerals in the podzol E horizon appeared to be perforated, as opposed to few if any in the abruptly underlying B horizon, suggesting a link to podzolization. High concentrations of Al and Si in organic surface horizons under boreal and temperate conifers can be explained by transfer by hypha of weathering products from the minerals to mycorrhizal roots in the O horizon, followed by release of weathering products that are not taken up by the plants. Rock-eating ectomycorrhizal fungi suggest a more direct role for plants in podzolization than hitherto realized, providing tight coupling between podzolization and mineral weathering. Preliminary observations, however, indicate that mycorrhizal fungi do not play a role in podzolization under Kauri ( Agathis australis) in New Zealand.

Mycorrhizal weathering: A true case of mineral plant nutrition?

Weatherable minerals in all podzol surface soils andshallow granitic rock under European coniferousforests studied hitherto are criss-crossed bynumerous open, tubular pores, 3–10 µm in width. Wehypothesize that these pores were formed bycomplex-forming, low-molecular weight organic acidsexuded by or formed in association with mycorrhizalfungi. It is well known that ectomycorrhizal myceliumrepresents a greatly extended, and better distributed,surface area for the absorption of nutrients. However, there have been few investigations of how thewhereabouts of individual hypha affect nutrientuptake. The results presented here provide directevidence that the mycelium is able to penetrate, andmost probably create, microsites which areinaccessible to plant roots and isolated from bulksoil solution phenomena. Dissolved products could betranslocated to the host plant roots, bypassing thesoil solution with often toxic concentration ofAl3+ from acid rain, and bypassing competitionfor nutrient uptake by other organisms. Furthermore,there is strong evidence that ``rock-eating''mycorrhizal fungi play a role in the formation ofpodzol E horizons. The partly speculativeinterpretations presented here challenge conventionalideas about (1) the importance of nutrient uptakefrom the bulk soil solution (2) criteria for criticalloads of acid atmospheric deposition for forests, and(3) the process of podzolization.