Bio—Mobilization of Potassium from Clay Minerals：II.By ectmycorrhizal Fungi

Ectomycorrhizal ungi,including Cenococcum geophilum SIV( Cg SIV),and Pisolithus tinctorius 2144(Pt 2144),441(Pt 441) and XC1(Pt XC1),were cultured in Pachlewski liquid medium with H2KPO4,KCl-saturated vermiculite and mica as K sources,respectively,to investigate the mechanism of K absorption and mobilization by the fungi,FUngal growth rate,K absoprtion and mobilization varied significantly among the fungal species,Faster growth and greater K accumulation in Pt XC1 than Pt 2144,Pt 441 and Cg siv were observed.Ectomycorrhizal fungi depressed HCl-soluble K in minerals after successive extractions by water and NH4OAc.Ratio of the total amount of K,including water-,NH4OAc-and HCl-soluble K,lost from substrates to the K accumulated in fungal colonies was less than 60%.These reveal that the ectomycorrhizal fungi could utilize K in Interlayer and structural pools,which are usually unavailable for plants in short period.Large differences in the depletion of K in interlayer and structural pools by fungi were observed at fungal harvest.Taking into account the nutrient absorption by ectomycorrhizal fungi in symbionts and the direct contact between hyphae and soils,the fungi species colonized on the root surfaces seemed to be related to the effectiveness of mycorrhizas to utilize K in soils.Ectomycorrhizal fungi differed in the efflux of protons and oxalate.Pt XC1 was observed to have greatest ability to effuse protons and oxalate among the fungi adopted in the experiment.Furthermore,the higther the concentrations of protons and oxalate in the liquid culture solutions,the larger the depletion of K in interlayer and structural pools in minerals by fungi,Protons could replace interlayer K and chelation of oxalate with Fe and Al in crystal lattice could cause weathering of clay minerals.So,protons and oxalate produced by ectomycorrhizal fungi might play an important role in K mobilization in these two pools.     

粘土矿物中钾的生物活化: Ⅱ. 外生菌根真菌(的作用)

Ectomycorrhizal fungi, including Cenococcum geophilum SIV (Cg SIV), and Pisolithus tinctorius 2144 (Pt 2144), 441 (Pt 441) and XC1 (Pt XC1), were cultured in Pachlewski liquid medium with H₂KPO₄, KCl-saturated vermiculite and mica as K sources, respectively, to investigate the mechanism of K absorption and mobilization by the fungi. Fungal growth rate, K absorption and mobilization varied significantly among the fungal species. Faster growth and greater K accumulation in Pt XC1 than Pt 2144, Pt 441 and Cg siv were observed. Ectomycorrhizal fungi depressed HCl-soluble K in minerals after successive extractions by water and NH₄OAc. Ratio of the total amount of K, including water-, NH₄OAc- and HCl-soluble K, lost from substrates to the K accumulated in fungal colonies was less than 60%. These reveal that the ectomycorrhizal fungi could utilize K in interlayer and structural pools, which are usually unavailable for plants in short period. Large differences in the depletion of K in interlayer and structural pools by fungi were observed at fungal harvest. Taking into account the nutrient absorption by ectomycorrhizal fungi in symbionts and the direct contact between hyphae and soils, the fungi species colonized on the root surfaces seemed to be related to the effectiveness of mycorrhizas to utilize K in soils. Ectomycorrhizal fungi differed in the efflux of protons and oxalate. Pt XC1 was observed to have greatest ability to effuse protons and oxalate among the fungi adopted in the experiment. Furthermore, the higher the concentrations of protons and oxalate in the liquid culture solutions, the larger the depletion of K in interlayer and structural pools in minerals by fungi. Protons could replace interlayer K and chelation of oxalate with Fe and Al in crystal lattice could cause weathering of clay minerals. So, protons and oxalate produced by ectomycorrhizal fungi might play an important role in K mobilization in these two pools.     

淹水后可变电荷土壤铁锰形态的转化

Ectomycorrhizal fungus Laccaria bicolor S238N,isolated from a forest soil in central France in 1990s,has demonstrated unequivocally and ability to promote pine growth.In the present nursery bed experiment,the ability of this ectomycorrhizal fungus to increase growth and P and K uptake of Douglas Fir seedlings (Zone 22) was examined.Growth of inoculated seedlings was over twice(plant height) and three times (biomass)that of non-inoculated ones.Similarly,both the concentrations and the amounts of P and K uptake by seedlings were significantly increased by fungal inoculation,indicating the improvement of P and K nutrition in mycorrhizal seedlings.In contrast,Al-P in the soils was decreased obviously by plants,especially by mycorrhizas,suggesting utilization of this soil P pool by plants and more efficient Al-P mobilization by mycorrhizas than by nomycorrhizas.Moreover,K extracted by 1mol/L HCl following consecutive extraction of H2O and CH3COONH4,which may not be plant available,could be utilized by fungus colonied roots.This could be explained by the release of protons and oxalate by hypae which leads to replacement of interlayer K in nonexpanded 2:1 clay minerals and bio-weathering of phyllosilicates.     

阳离子交换膜在土壤钾释放及小麦对钾素的反应研究中的应用

A pot experiment was conducted in the growth chamber on Saskatchewan soils with different texture to determine the K release status and wheat K demand.The relationship between K uptake and soil available K extracted by cation exchange membrane(CEM-K) and the effcet of K fertilizer on wheat growth and soil available K was also evaluated.Treatments of 0,60 and 120mg K/kg were applied to sandy,low and high K loamy and clay soils,The highest yields were acieved with the application of 120mg K/kg in sandy soil and 60 mg K/kg in other soils.On the whole,the clay soil contributed K more than other soils from slowly available fraction.Regression revealed a linear relationship between the soil available K extracted by NH4OAc(Ka) and CEM-K in suspensions(r=0.93).Results also showed that CEM-K in burial and in suspensions were different not only in the amount but also in correlation with Ka or K uptake.     

中国福建省土壤的主要粘土矿物

The clay minerals of more than 200 soil samples collected from various sites of Fujian Province were studied by the X-ray diffraction method and transmission electron microscopy to study their distribution and evolution.Montmorillonite was found in coastal solonchak,paddy soils derived from marine deposit,lacustrine deposit and river deposit,and some lateritic red soil,red soil and yellow soil with a low weathering degree.Chlorite existed mainly in coastal solonchak and paddy soil developed from marine deposit.1.4nm intergradient mineral appeared frequently in yellow soil,red soil and lateritic red soil.The content of 1.4nm intergradient mineral increased with the decrease of weathering degree from lateritic red soil to red soil to yellow soil.Hydrous micas were more in coastal solonchak,paddy soils derived from marine deposit,lacustrine deposit and river deposit.and puple soil from purple shale than in other soils.Kaolinte was the most important clay mineral in the soils iun this province.The higher the soil weathering degree,the more the kaolinite existed.From yellow soil to red soil to lateritic red soil,kaolinite increased gradually,Kaolinite was the predominant clay mineral accompanied by few other minerals in typical lateritic red soil. Tubular halloysite was a widespread clay mineral in soils of Fujian Province with varying quantities.The soil derived from the paent rocks rich in feldspar contained more tubular halloysite.Spheroidal halloysite was found in a red soil and a paddy soil developed from olivine basalt gibbsite in the soils in this district was largely“primary gibbsite” which formed in the early weathering stage.Gibbsite decreased with the increase of weathering degree from yellow soil to red soil to lateritic red soil.Goethite also decreased in the same sequence while hematite increased.     

外生菌根真菌对土壤无机磷迁移的影响

Ectomycorrhizal(EM) fungi could form symbiosis with plant roots and participate in nutrient absorption; however, many EM species commonly found in forest soils, where phosphorus(P) concentration and availability are usually very low, particularly in tropical and subtropical areas, have not yet been investigated for their efficiencies to mobilize soil P. In this study, fungal growth, P absorption,efflux of protons and organic acids, and soil P depletion by four isolates of EM fungi isolated either from acidic or calcareous soils were compared in pure liquid culture using soil as a sole P source. Boletus sp. 7(Bo 7), Lactarius deliciosus 3(Ld 3), and Pisolithus tinctorius 715(Pt 715) from acidic and P-deficient soils of southwestern China showed higher biomass and P concentration and accumulation than Cenococcum geophilum 4(Cg 4) from a calcareous soil of Inner Mongolia, northern China, after 4 weeks of liquid culture. Oxalate, malate, succinate, acetate, and citrate concentrations in the culture solutions varied significantly with fungal species,and oxalate accounted for 51.5%–91.4% of the total organic acids. Organic acids, particularly oxalate, in the culture solutions may lead to the solubilization of iron-bound P(Fe-P), aluminum-bound P(Al-P), and occluded P(O-P) from soil phosphates. Fungal species also varied greatly in proton efflux, which decreased the culture solution pH and may dissolve calcium-bound P(Ca-P) in soil.This could be the reason for the increment of both inorganic P in the culture solutions and Olsen P in the soil when EM fungi were present. Total inorganic P, the sum of Al-P, Fe-P, O-P, and Ca-P, in the culture solutions was positively correlated with the total concentration of organic acids in the culture solutions(r = 0.918*, n = 5), but negatively with both the total inorganic P in soil(r =-0.970**, n = 5) and the culture solution pH(r =-0.830*, n = 5). These suggested variable efficiencies of EM fungal species to mobilize inorganic P fractions from soil, which could make EM trees to utilize inorganic P in the same way like EM fungi and adapt to the soils with various P concentrations and availabilities.     

应用反射光谱法对土壤粘土矿物的鉴定的初步研究

The characteristics of the reflectance spectra of clay minerals and their influences on the reflectance spectra of soils are dealt with in the paper.The results showed that dominant clay minerals in soils could be distinguished in light of the spectral -form parameters of the reflectance spectra of soils,thus making it possible to develop a quick method to determine clay minerals by means of reflectance spectra of soils in the lab.and providing a theoretic basis for remote sensing of clay minerals in soils with a high resolution imaging spectrometer.     

土壤非交换态钾与结构态钾能够区分吗?（英文）

Nonexchangeable K(NEK) is the major portion of the reserve of available K in soil and a primary factor in determining soil K fertility.The questions of how much NEK is in soils and how to quantify total NEK in soils are so far still unclear due to the complicated effects of various minerals on K fixation.In this study,the NEK in 9 soils was extracted with sodium tetraphenylboron(NaBPh_4) for various time periods longer than 1 d.The results showed that the NEK extracted by NaBPh4 gradually increased with time,but showed no more increase after the duration of extraction exceeded 10-20 d.As the temperature increased from 25 to 45 ℃,the duration to obtain the maximum extraction of NEK was reduced from 20 to 10 d,and the maximum values of NEK released at both temperatures was almost the same for each soil.The maximum NEK(MNEK) of the 9 soils extracted by NaBPh.4 varied from3074 to 10081 mg kg~(-1),accounting for 21%-56%of the total soil K.There was no significant correlation between MNEK released by NaBPh4 and other forms of K,such as NH_4OAc-extracted K,HNOg-extracted K and total K in soils,which indicates that NEK is a special form of K that has no inevitable relationship to the other forms of K in soils.The MNEK extraction by NaBPh4 in this study indicated that the total NEK in the soils could be differentiated from soil structural K and quantified with the modified NaBPlu method.The high MNEK in soils made NEK much more important in the role of the plant-available K pool.How to fractionate NEK into different fractions and establish the methods to quantify each NEK fraction according to their bioavailability is of great importance for future research.     

外生菌根诱导油松根际土壤重金属形态变化

To understand the role of ectomycorrhizas in improving the tolerance of its host to excessive heavy metals in soil, this study was conducted to exam the patterns of four fractions （the exchangeable, the carbonate-bound, the Fe-Mn oxide- bound and the organically bound） of both Cu and Cd in the rhizosphere of Chinese pine （Pinus tabulaeformis） seedlings grown in excessive Cu and Cd environment. The results showed that the speciation of Cu and Cd in the rhizosphere was significantly influenced by inoculation of ectomycorrhizal fungus Boletus edulis. Compared to the rhizosphere, the content of exchangeable Cu slightly decreased in the mycorrhizosphere of the seedlings grown in 166 and 400 mg kg^-1 Cu contaminated soil, whereas the exchangeable Cd in the mycorrhizosphere decreased remarkably to only 33% and to 60% that of the rhizosphere at 0.75 and 1.50 mg kg^-1 Cd levels, respectively. These indicate the potential capacity of mycorrhizas to alleviate the damage of heavy metals to the host plants by reducing the bioavailability of heavy metals in soil. Distribution of the 4 tested fractions of Cu and Cd at different contamination levels showed that there was a strong tendency of changing from loosely associated fractions to strongly associated fractions in the mycorrhizosphere. The most stable Cd fraction, organically bound Cd, was significantly larger in the mycorrhizosphere than in the rhizosphere at different Cd contamination levels. This phenomenon was also observed for Cu but the difference was not statistically significant.     

草酸对土壤和矿物中钾素的释放规律研究

Oxalic acid plays an important role in improving the bioavailability of soil nutrients. Batch experiments were employed to examine the influences of oxalic acid on extraction and release kinetics of potassium （K） from soils and minerals along with the adsorption and desorption of soil K^＋. The soils and minerals used were three typical Chinese soils, black soil （Mollisol）, red soil （Ultisol）, and calcareous alluvial soil （Entisol）, and four K-bearing minerals, biotite, phlogopite, muscovite, and microcline. The results showed that soil K extracted using 0.2 mol L^-1 oxalic acid was similar to that using 1 mol L^-1 boiling HNO3. The relation between K release （y） and concentrations of oxalic acid （c） could be best described logarithmically as y = a ＋ blogc, while the best-fit kinetic equation of K release was y = a ＋ b√t, where a and b are the constants and t is the elapsed time. The K release for minerals was ranked as biotite 〉 phlogopite 〉〉 muscovite 〉 microcline and for soils it was in the order： black soil 〉 calcareous alluvial soil 〉 red soil. An oxalic acid solution with low pH was able to release more K from weathered minerals and alkaline soils. Oxalic acid decreased the soil K^＋ adsorption and increased the soil K^＋ desorption, the effect of which tended to be greater at lower solution pH, especially in the red soil.     

Ectomycorrhizal Association Enhances Al Tolerance by Inducing Citrate Secretion in Pinus densiflora

To investigate the effects of ectomycorrhizal association on the aluminum (Al) tolerance of Pinus densiflora Sieb. & Zucc., seedlings with or without ectomycorrhizal association with the fungus Pisolithus tinctorius (Pers.) Coker & Couch were exposed to 1 mM Al. Association with P. tinctorius alleviated Al-induced inhibition of root elongation and biomass growth in the mycorrhizal seedlings. Secretion of malate and citrate, both low-molecular-weight organic acids that could detoxify Al by the formation of stable complexes, was investigated in P. tinctorius mycelia and in pine roots with and without P. tinctorius association. Citrate secretion from the P. tinctorius mycelia in vitro was stimulated by Al. Citrate secretion from the roots of the ectomycorrhizal seedlings was also stimulated by Al, but was not detected in the non-mycorrhizal seedlings. These results suggest that citrate secreted from the roots of the ectomycorrhizal seedlings was produced in the hyphae of P. tinctorius. Citrate secretion may play a role in enhancing the Al tolerance of host seedlings.     

Release of interlayer potassium in Norwegian grassland soils

Potassium (K) release from sources that are not initially exchangeable is attributed to depletion of interlayer K of micas and clay minerals or weathering of feldspars. The aim of the present study was to estimate the K release from interlayer K. Soil samples from 17 field experiments in ley on a range of mineral soils in Norway were used in the study. The change in K‐fixation capacity was used as an estimate of depletion of interlayer K. It was assumed that the increase in K‐fixation capacity during 3 yr of cropping was equivalent to the amount of K depleted from the interlayer positions. Mean K fixation increased in the majority of the soils during 3 yr of grass cropping both with and without K application. The increase in K fixation indicated that without K application, the K uptake from interlayer K amounted to 43%, 28%, and 26% of the K yield for clay soils, high‐K sandy soils, and low‐K sandy soils, respectively. Including K uptake from exchangeable K in the topsoil and from subsoil, the explained K uptake amounted to 79%, 69%, and 81% for the three groups of soil, respectively. Simple linear‐regression analyses showed that the change in K fixation during 3 yr of grass cropping was best explained by the percentage of clay in the soil.     

山东主要果园土壤的粘土矿物组成及其吸附特性

研究了山东主要果园土壤中粘土矿物的组成、类型及其对P、K和Cu、Zn、Pb金属元素的吸附特性。结果表明：山东主要果园土壤的粘土矿物类型存在着明显的差异,淋溶较强,酸度较大的棕壤（简育湿润淋溶土）中粘土矿物以高岭石占优势,对P的吸附较强,但对K＋吸附固定较弱。含游离C aCO3较高的潮土（淡色潮湿雏形土）和褐土（简育干润淋溶土）,对P有较强的吸附和沉淀作用,使土壤磷的有效性降低。而砂姜黑土（钙积潮湿变性土）由于含有较高的蒙脱石和1.4 nm过渡矿物,对K＋具有很强的吸附和晶穴固定作用,因此砂姜黑土中磷肥和钾肥的有效性均较低,在施肥上应采取集中施肥和保持较湿润土壤环境等措施,以提高养分有效性。砂姜黑土和潮土对Cu、Zn、Pb金属元素的吸附显著地大于棕壤和褐土,主要的影响因素是不同土壤的粘粒含量和粘土矿物的类型的差异。同一土壤对Pb的吸附量远远大于对Zn和Cu的吸附量,主要取决于金属元素本身的化学性质和胶体的吸附特性。     

Rapid incorporation of carbon from ectomycorrhizal mycelial necromass into soil fungal communities

Ectomycorrhizal mycelial necromass is an important source of carbon for free-living microorganisms in forest soils, yet we know little either of its fate when it enters soil or of the identity of microbes that are able to utilise mycelium as their energy source. Here we used ¹³C-labelled mycelium of the ectomycorrhizal fungus Pisolithus microcarpus in laboratory incubations in combination with DNA-stable isotope probing (SIP) to determine the identity of functionally active soil fungi that can utilise dead mycelium. We also used solid-state nuclear magnetic resonance (NMR) spectroscopy to detect parallel changes in the abundance of key biochemical constituents of soil. A decrease in bulk soil ¹³C concentration together with rapid loss of glycogen and chitin-glucan during the 4 week incubations suggested that dead mycelium was rapidly turned over. Further, ¹³C was incorporated into fungal DNA within 7 days of addition to soil. DNA-SIP also revealed a dynamic community of functionally active soil fungi. By applying DNA-SIP and NMR in parallel, our data show that carbon from decaying ectomycorrhizal mycelium is rapidly transformed and incorporated into free-living soil fungi. This finding emphasises that dead extra-matrical mycelium is an important source of labile carbon for soil microorganisms.     

Bedeutung der Kalium-Verarmung in der Rhizosphäre und der Tonminerale für die Freisetzung von nichtaustauschbarem Kalium und dessen Bestimmung mit HCl

Significance of K-depletion in the rhizosphere and clay minerals for the release of nonexchangeable potassium and its determination with HCl Experiments were carried out with four different soils (basaltic soil, C-horizon of a keuper, sea alluvium, and a podsolic brown earth from loess) to investigate whether the fractions of exchangeable K and HCl-soluble nonexchangeable K are decreased in root vicinity by the K uptake of ryegrass (Lolium perenne). After a growth period of 70 days with five cuts distinct K depletion profiles were obtained showing that in the direct vicinity of the root the decrease in exchangeable K was the most pronounced. The K depletion in this fraction extended to 2 cm distance from the root zone. Also the concentration of HCl-soluble nonexchangeable K of the interlayers was decreased in root vicinity. It is assumed that the decrease in K concentration in soil solution is a prerequisite for the net release of interlayer K. The decrease in exchangeable K plus HCl soluble nonexchangeable K was equal to the amount of K taken up by the grass from the basaltic- and the alluvial soil. In the case of the podsolic brown earth and the keuper soil, however, much more K was taken up by the grass than was found in the decrease of exchangeable K plus HCl soluble nonexchangeable K. It is assumed that in these two soils the fraction of the HCl soluble K was more altered by the expansion of interlayers during the growth of rye-grass. The net release of interlayer K was neither related to the concentration of exchangeable K, HCl-soluble non-exchangeable K, the K fixation power nor the clay concentration. It is supposed that the net release of interlayer K also depends on clay mineral characteristics.     

Reforestation of burned stands: The effect of ectomycorrhizal fungi on Pinus pinaster establishment

The area occupied by Pinus pinaster in Portugal is rapidly diminishing because of forest fires. Ectomycorrhizal fungi form obligate, mutually beneficial associations with P. pinaster which improve plant growth and resistance to adverse conditions. The aim of this work was to assess whether native ectomycorrhizal fungi could be a useful tool in the reforestation of burned areas. The work was conducted in a forest nursery greenhouse, where P. pinaster seedlings were inoculated with compatible ectomycorrhizal fungal isolates: Suillus bovinus, Pisolithus tinctorius, Rhizopogon roseolus, and a mixture of the three fungi, using burned and unburned forest soil as substrate. Inoculation significantly enhanced the growth of P. pinaster, with R. roseolus proving to be the most effective in burned soil, with an 8-fold increase in plant fresh weight. Overall, inoculation stimulated growth most in burned than in unburned soil.This study suggests that inoculation with selected ectomycorrhizal fungi in containerised nurseries can be an advantageous approach for the successful establishment of P. pinaster in burned soil. The obtained results point out to the interest of extending these studies into fire-impacted areas, using ectomycorrhizal fungi as a biological tool.     

Mobilization of aluminium, iron and silicon by Picea abies and ectomycorrhizas in a forest soil

Weathering of soil minerals is a key determinant of ground and surface water quality and is also important in pedogenic and rhizosphere processes. The relative importance of biotic and abiotic studies in mineral weathering, however, is poorly understood. We investigated the impact of Picea abies seedlings, an ectomycorrhizal fungus and humic acid on the solubilization of aluminium (Al), iron (Fe) and silicon (Si) in an E horizon forest soil over 10 months. Elemental budgets were constructed based upon losses in drainage water, accumulation in plants and changes in the pools of exchangeable ions. Plants and mycorrhizas or both had a significant effect on the total amounts of Al, Fe and Si mobilized from the soil. Significantly larger amounts of Al and Fe were recovered in plants than those lost in drainage water, whereas the opposite trend was true for Si. The continual addition of dissolved organic matter to the soil in the form of humic acid had an effect only on mobilization of Fe, which increased due to larger plant uptake and an increase in the exchangeable pool. The mobilization of Fe and Si were positively correlated with hyphal length, soil respiration and concentrations of oxalate in the soil solution, and mobilization of Al was strongly correlated with plant weight. Scanning electron microscopy revealed that most fungal hyphae were associated with mineral surfaces with little occupation of cracks and micropores within mineral grains. Evidently ectomycorrhizas have important impacts on mineral dissolution and the chemistry of forest soils.