Effects of phosphoroamides on ammonia volatilization and nitrite accumulation in soils treated with urea

Summary We compared the effects of N-(n-butyl) thiophosphoric triamide (NBPT), N-(diaminophosphinyl)-cyclohexylamine (DPCA), phenylphosphorodiamidate (PPD), and hydroquinone on transformations of urea N in soils. The ability of these urease inhibitors to retard urea hydrolysis, ammonia volatilization, and nitrite accumulation in soils treated with urea-decreased in the order NBPT > DPCA PPD > HQ. When five soils were incubated at 30°C for 14 days after treatment with urea (1 mg urea N g^–1 soil), on average, the gaseous loss of urea N as ammonia and the accumulation of urea N as nitrite were decreased from 52 to 5 % and from 11 to 1%, respectively, by addition of NBPT at the rate of 10 g g^–1 soil (0.47 parts of NBPT per 100 parts of urea). The data obtained support previous evidence that NBPT is more effective than PPD for reduction of the problems encountered in using urea as a fertilizer and deserves consideration as a fertilizer amendment for retarding hydrolysis of urea fertilizer in soil.     

Movement of urea and its hydrolysis products as influenced by moisture content and urease inhibitors

A laboratory experiment was conducted on an Aquic Udifluvent Belgian soil in order to study the movement of urea and its hydrolysis products. This study was carried out at two moisture levels (10 and 20%) upon the addition of three types of urease inhibitors: hydroquinone, phenylphosphorodiamidate (PPDA), and N-(n-butyl)phosphorothioic triamide (NBPT). The results clearly show the effects of the inhibitors in retarding the hydrolysis of urea. The highest effect was observed with NBPT, followed by hydroquinone, and PPDA. The effect was more pronounced at 10% than at 20% moisture content. It was clear that subsequent nitrification of the NH _inf4 ^sup+ formed was inhibited at the lower moisture level. At 10% moisture, from the 7th day of incubation on, some NH _inf4 ^sup+ moved about 3 cm and reached the top of the soil column. At 20% moisture, no NH _inf4 ^sup+ reached the surface as it was quickly nitrified. After 17 days of incubation and at 20% moisture, the total mineral N was more or less homogeneously distributed within the soil column. In contrast, at 10% moisture, the remaining urea and the hydrolysis products were still concentrated at the place of application. The distribution of urea and its hydrolysis products was comparable with 7 days of incubation at 20% moisture and 17 days at 10%.     

Effect of N-(n-butyl) thiophosphoric triamide on hydrolysis of urea by plant,microbial, and soil urease

Summary Comparison of the effects of N-(n-butyl) thiophosphoric triamide (NBPT) and phenylphosphorodiamidate (PPD) on hydrolysis of urea by plant (jackbean), microbial (Bacillus pasteurii), and soil urease showed that whereas NBPT was considerably more effective than PPD for inhibiting hydrolysis of urea added to soil, it was much less effective than PPD for inhibiting hydrolysis of urea by plant or microbial urease. Studies to account for this observation indicated that NBPT is rapidly decomposed in soil to a compound that is much more effective than NBPT for inhibition of urease activity and that this compound is N-(n-butyl) phosphoric triamide.     

Urea N uptake efficiency of ryegrass (Lolium perenne L.) in the presence of urease inhibitors

Summary A greenhouse experiment was conducted to study the comparative efficiency of urea as an N fertilizer with and without the addition of different urease inhibitors. Ryegrass (Lolium perenne L.) was used as the test plant and the N balance technique with ¹⁵N was applied. Three urease inhibitors, hydroquinone, phenyl phosphorodiamidate (PPDA), and N-(n-butyl) phosphorothioic triamide (NBPT), were evaluated for their effects on urea-N uptake as well as on grass yield. The addition of urease inhibitors, except for hydroquinone in the later growth period, did not significantly influence the dry matter weight. Throughout the whole growth period, only NBPT significantly increased the total urea-N uptake. In the uninhibited system, the major fertilizer N loss occurred during the first period of grass growth, presumably via NH₃ volatilization, since the environment did not favour the other pathways of N loss. However, an appreciable amount of urea N was lost during the later growth period in all inhibited systems, especially in the hydroquinone-treated system. This indicates that the application of urease inhibitors could not eliminate the urea N loss. The greater N loss in the hydroquinone-treated soil appears to be related to the inhibition by hydroquinone of nitrification.     

Effects of the urease inhibitor N-(n-butyl)phosphorothioic triamide in low concentrations on ammonia volatilization and evolution of mineral nitrogen

Laboratory incubation experiments were conducted to study the influence of increasing concentrations of N-(n-butyl)phosphorothioic triamide (NBPT) on NH₃ volatilization and rate of urea hydrolysis and evolution of mineral N in Ozzano, Rimini and Carpi soils with different physicochemical characteristics. Low concentrations of NBPT reduced NH₃ losses due to volatilization after urea fertilization and the effectiveness of the inhibitor was related to the soil characteristics (e.g. high concentrations of organic C and sand). After 15 days of incubation, no significant reductions of losses were found for any of the NBPT concentrations employed in Rimini soil. The application of NBPT led to a considerable reduction of the formation of nitrite. This process was completely annulled with the highest dose of NBPT (0.5% w/w_urea) in the Carpi soil after 15 days. In Rimini soil, however, the use of NBPT was less effective in influencing nitrite formation. The use of NBPT favoured accumulation of nitrate proportional to the NBPT concentration employed while it had no influence on the NH _inf4 ^sup+ fixation by 2:1 layer silicates. The data obtained support previous evidence that NBPT is effective in reducing the problems encountered in using urea as fertilizer. However, environmental conditions and soil physicochemical characteristics may have an important influence on the effectiveness of NBPT.     

Evaluation of 3-methylpyrazole-1-carboxamide as a soil nitrification inhibitor

Summary Laboratory studies to evaluate 3-methylpyrazole-1-carboxamide (MPC) as a soil nitrification inhibitor showed that it was comparable to nitrapyrin (N-Serve) for inhibiting nitrification of ammonium in soil, but was not as effective as etridiazole (Dwell) or 2-ethynylpyridine. They also showed that the effectiveness of MPC as a soil nitrification inhibitor is markedly affected by soil type and soil temperature, that MPC is more effective for inhibiting nitrification of ammonium-N than of urea-N, and that MPC has little, if any, effect on hydrolysis of urea or denitrification of nitrate in soil. These observations and other work discussed indicate that MPC is one of the most promising compounds so far proposed for inhibition of nitrification in soil.     

Effect of urease inhibitors on urea hydrolysis and ammonia volatilization

Summary Two laboratory incubation experiments were conducted to study the effects of the urease inhibitors hydroquinone (HQ), phenyl phosphorodiamidate (PPDA), and N-(n-butyl) thiophosphoric triamide (NBPT) in retarding the hydrolysis of urea, in the evolution of mineral N, and in reducing NH₃ loss through volatilization, under aerobic and waterlogged conditions, both at 25°C. NBPT generally exceeded PPDA and HQ in the ability to delay urea hydrolysis and NH _inf4 ^sup+ accumulation under aerobic conditions, whereas PPDA retarded these activities more effectively under anaerobic conditions. HQ was less effective than the other two urease inhibitors. Under aerobic conditions, 20% of the applied urea was lost through NH₃ volatilization after 5 days in the system without an inhibitor. With the addition of HQ and PPDA, the volatilization was delayed by 1 day but not eliminated. NBPT effectively decreased the NH₃ loss, from 20 to 3% of the applied urea. A more severe N loss (40%) occurred in the waterlogged system. HQ had little effect on NH₃ volatilization. PPDA decreased the NH₃ loss from 40 to less than 20% of the applied urea. The effectiveness of NBPT decreased under anaerobic conditions. It was concluded that urease inhibitors can reduce NH₃ volatilization following the application of urea. However, environmental conditions might have an important influence on the effectiveness of these inhibitors.     

N-丁基硫代磷酰三胺对淹水土壤脲酶动力学特征的影响

研究了淹水状态下脲酶抑制剂N-丁基硫代磷酰三胺(NBPT)对东北4种典型土壤(黑土、白浆土、棕壤、褐土)脲酶动力学参数的影响。结果表明,NBPT对土壤脲酶动力学参数的作用程度受土壤类型、培养时间及其交互作用的显著影响。与对照相比,NBPT使土壤脲酶米氏常数(Km)增加,最大反应速率(Vmax)和Vmax/Km降低,表明NBPT对土壤脲酶的作用机理属于典型的混合型抑制。随培养时间延长,土壤脲酶Km降低,Vmax和Vmax/Km升高,表明NBPT对脲酶的抑制效力逐渐减弱。黑土、白浆土脲酶Km值小于棕壤和褐土,而Vmax则大于棕壤和褐土,表明土壤肥力水平能够影响土壤脲酶的动力学参数。相关分析表明,土壤脲酶Km与粘粒和粉粒含量呈显著正相关关系,与pH值呈显著负相关关系;而Vmax和Vmax/Km的相关性则正好与此相反。     

Suppression of methane oxidation in aerobic soil by nitrogen fertilizers,nitrification inhibitors,and urease inhibitors

Concentrations of CH₄, a potent greenhouse gas, have been increasing in the atmosphere at the rate of 1% per year. The objective of these laboratory studies was to measure the effect of different forms of inorganic N and various N-transformation inhibitors on CH₄ oxidation in soil. NH ₄ ⁺ oxidation was also measured in the presence of the inhibitors to determine whether they had differential activity with respect to CH₄ and NH ₄ ⁺ oxidation. The addition of NH₄Cl at 25 g N g^-1 soil strongly inhibited (78–89%) CH₄ oxidation in the surface layer (0–15 cm) of a fine sandy loam and a sandy clay loam (native shortgrass prairie soils). The nitrification inhibitor nitrapyrin (5 g g^-1 soil) inhibited CH₄ oxidation as effectively as did NH₄Cl in the fine sandy loam (82–89%), but less effectively in the sandy clay loam (52–66%). Acetylene (5 mol mol^-1 in soil headspace) had a strong (76–100%) inhibitory effect on CH₄ consumption in both soils. The phosphoroamide (urease inhibitor) N-(n-butyl) thiophosphoric triamide (NBPT) showed strong inhibition of CH₄ consumption at 25 g g^-1 soil in the fine sandy loam (83%) in the sandy clay loam (60%), but NH ₄ ⁺ oxidation inhibition was weak in both soils (13–17%). The discovery that the urease inhibitor NBPT inhibits CH₄ oxidation was unexpected, and the mechanism involved is unknown.     

土壤温度和含水量互作对抑制剂抑制氮素转化效果的影响

为比较生化抑制剂组合对土壤氮素转化的抑制效果,揭示不同土壤温度和含水量互作对尿素水解抑制效应的影响。该文采用室内模拟培养方法,研究土壤含水量(60%和80%田间最大持水量,water holding capacity,WHC)和土壤温度(15、25和35℃)互作对生化抑制组合[N-丁基硫代磷酰三胺(N-(n-butyl)thiophosphoric triamide,NBPT)、N-丙基硫代磷酰三胺(N-(n-propyl)thiophosphoric triamide,NPPT)和2-氯-6(三氯甲基)吡啶(2-chloro-6(trichloromethyl)pyridine,CP)在黄泥田土壤中抑制氮素转化效果的影响。结果表明:土壤温度和含水量对生化抑制组合在黄泥田土壤中抑制尿素水解效应显著,以土壤温度影响更大。随着土壤温度增加,尿素水解转化增强,有效作用时间降低,硝化作用增强,脲酶和硝化抑制效应减弱;随着土壤含水量降低,尿素水解转化缓慢,有效作用时间延长,硝化作用减弱,脲酶和硝化抑制效应增强。不同土壤温度和含水量条件下,NBPT/NPPT或配施CP处理有效抑制黄泥田土壤脲酶活性,延缓尿素水解;CP或配施NBPT/NPPT处理有效抑制NH4+-N向NO_3~--N转化,保持土壤中较高NH_4~+-N含量长时间存在。新型脲酶抑制剂NPPT单独施用及与CP配施的土壤尿素水解抑制效果与NBPT相似。黄泥田土壤中生化抑制组合应用最佳的土壤温度和含水量分别为25℃和60%WHC。总之,针对不同土壤温度和含水量条件,在黄泥田土壤中应采用脲酶抑制剂与硝化抑制剂相结合的施肥方式。     

Effect of organic matter and urease inhibitors on urea hydrolysis and immobilization of urea nitrogen in an alkaline soil

Summary Laboratory incubation experiments with ¹⁵N-labelled urea were conducted on a Aquic Udifluvent Belgian soil amended with barley straw, in, order to study the influence of three urease inhibitors, hydroquinone, phenyl phosphorodiamidate and N-(n-butyl) phosphorothioic triamide on urea hydrolysis and N transformations. The results demonstrated that the urea was hydrolyzed more rapidly when the soil was amended with ground barley straw. A pronounced inhibition of urease inhibitors occurred with the urea hydrolysis, but it was decreased by increasing the soil organic C content. A severe N immobilization (about 82–100% of the applied urea) occurred in soil samples that were rich in organic C. The addition of urease inhibitors increased urea-N immobilization by 5–30%. N-(n-butyl) phosphorothioic triamide had a stronger effect than the other two inhibitors when they were applied at the rate of 1%. However, the inhibitors decreased N immobilization when the soil was amended with barley straw.     

Use of palm-mat geotextiles for soil conservation: I. Effects on soil properties

Despite geotextile-mats having the potential for soil conservation, field studies on the effects of geotextiles on soil properties are limited. Hence, the utilization of palm-mat geotextiles as a potential soil conservation technique was investigated at Hilton, east Shropshire, U.K. (52°33′5.7″ N, 2°19′18.3″ W). Geotextile-mats constructed from Borassus aethiopum (Borassus palm of West Africa) and Mauritia flexuosa (Buriti palm of South America) leaves are termed Borassus mats and Buriti mats, respectively. Field experiments were conducted at Hilton during 2007–2009, to study the impacts of Borassus and Buriti mats on selected properties of the topsoil (0–5 cm). Ten fixed plots (10 × 1 m on a 15° slope) were established, with duplicate treatments. The treatments were: (i) bare soil; (ii) permanent grassed; (iii) bare soil with 1 m Borassus-mat buffer strips (area coverage ~ 10%) at the lower end of the plots; (iv) bare soil with 1 m Buriti mat buffer strips (area coverage ~ 10%) at the lower end of the plots; and (v) completely-covered with Borassus mats. Initial and final soil samples of the topsoil were collected and analysed for bulk density, aggregate stability, soil organic matter (SOM), total soil C (TSC), total soil N (TSN) and pH. Results indicate that, apart from Borassus completely-covered plots, soil bulk density increased and aggregate stability decreased in all plots after two years. Despite decreases in SOM contents in bare plots, SOM content did not change after two years in the grassed and geotextile treated plots. Treatments had no effects on changes in pH, TSC or TSN. Both Borassus and Buriti mat-covers within the buffer strip plots had little impact on SOM, TSC and TSN changes compared with bare soils within the same plots. Thus, Borassus buffer strip plots were very effective in maintaining some soil properties (i.e. SOM, TSC, and TSN) after two years of erosion by water. In summary, utilization of Borassus mats as buffer strips was very successful in conserving soil properties on a loamy sand soil.     

Effect of methods of application on the efficiency of urea broadcast onto lowland rice (Oryza sativa L.)

Summary We evaluated the effect of different methods of application on the efficiency of urea broadcast at a rate of 100 kg N ha^-1 onto lowland rice (Oryza sativa L. var. SPR 60) in a field experiment conducted on a Phimai soil (Fluvic Tropaquepts) during the dry season of 1989. Analysis of the floodwater on the first day after the fertilizer application showed a high initial concentration of urea-N. Addition of the urease inhibitor N-(n-butyl) thiophosphoric triamide (nBTPT), broadcast with the urea into the floodwater, caused an apparent reduction in the rate of urea disappearance and a subsequent accumulation of NH₃–N in the floodwater; this ureas inhibitor also suppressed the rise in floodwater pH, with a resultant reduction in the partial pressure of ammonia (pNH₃) compared with the unamended urea application. The use of nBTPT did not decrease the N loss from broadcast urea not did it increase the grain yield. Among the different methods of applying broadcast urea that we tested, the broadcast application of granular urea onto drained soil shortly after removing floodwater followed by flooding 2 days later appeared to be a good N management practice, offering considerable potential for improving the efficiency of urea applied to lowland rice crops.     

Effects of legume green-manuring on nitrogen mineralization and some microbiological properties in an acid rice soil

We studied N mineralization of legume green manures under laboratory and field conditions, and the effects of field green-manuring on the microbiological properties of an acid Alfisol soil. No significant differences were found in the mineralization rates of Sesbania (Sesbania cannabina), sunnhemp (Crotalaria juncea), and cowpea (Vigna unguiculata) green manure. Mineralization was higher in field-capacity moist soil than water-saturated soil. The decomposition of sunnhemp under field wetland conditions, in the absence of a rice crop, was a rapid as it was under in vitro conditions. The decomposition released considerable amounts of mineral N and the level of NH ₄ ⁺ -N was significantly higher than NO ₃ ^– -N. Significant improvements were observed in the microbial biomass, dehydrogenase activity, and bacterial populations in the field soil green-manured for rice for 3 years, compared with fertilized soil.     

Effect of nitrogen application on the A N value of soil

Pot experiments were conducted with two soils, from Rottenhaus and Seibersdorf in Austria, to ascertain whether the rate of fertilizer N application and the test crop would influence the amount of N available in the soil as assessed by the A-value method. ¹⁵N-labelled fertilizer was applied at rates of 10, 25, 40, 60, and 100 mg N kg^-1 soil, corresponding approximately to 20, 50, 80, 120 and 200 kg N ha^-1 respectively, and two crop species, barley (Hordeum vulgareL.) and non-nodulating soybean (Glycine max L.) were used to determine the soil A _N value under the various fertilizer regimes. The results showed that the Rottenhaus soil had a higher A _N value than the Seibersdorf soil, suggesting that the former was more fertile than the latter. The A _N values of both soils were significantly affected by the level of N application. When grown in the same soil, the two test crops showed significantly different fertilizer use efficiency and per cent N derived from fertilizer when the rate of N application exceeded 20 kg ha^-1. Thus, the A _N value as determined by the two test crops differed significantly for the same soil when the rate of N application was greater than 20 kg/ha. The difference was greater when the soil fertility level was high. The dependence of the A _N value on the level of N application and the species of crop seriously compromises the suitability of this method for determining plant-associated N₂ fixation. Hence, considerable caution is required when using this method to estimate plant-associated N₂ fixation.     

Trace gas emissions from soil of the central highlands of Mexico as affected by natural vegetation: a laboratory study

In the central highlands of Mexico, mesquite (Prosopis laevigata) and huisache (Acacia schaffneri), N₂-fixing trees or shrubs, dominate the vegetation and are currently used in a reforestation program to prevent erosion. We investigated how natural vegetation or cultivation of soil affected oxidation of CH₄, and production of N₂O. Soil was sampled under the canopy of mesquite (MES treatment) and huisache trees (HUI treatment), outside their canopy (OUT treatment) and from fields cultivated with maize (ARA treatment) at three different sites while production of CO₂, and dynamics of CH₄, N₂O and inorganic N (NH₄⁺, and NO₃^–) were monitored in an aerobic incubation. The production of CO₂ was 2.3 times higher and significantly greater in the OUT treatment, 3.0 times higher in the MES treatment and 4.0 times higher in the HUI treatment compared to the ARA treatment. There was no significant difference in oxidation of CH₄ between the treatments, which ranged from 0.019 g CH₄–C kg^–1 day^–1 for the HUI treatment to 0.033 CH₄–C kg^–1 day^–1 for the MES treatment. The production of N₂O was 30 g N₂O–N kg^–1 day^–1 in the MES treatment and >8 times higher compared to the other treatments. The average concentration of NO₃^– was 2 times higher and significantly greater in the MES treatment than in the HUI treatment, 3 times greater than in the OUT treatment and 10 times greater than in the ARA treatment. It was found that cultivation of soil decreased soil organic matter content, C and N mineralization, but not oxidation of CH₄ or production of N₂O.     

土壤团聚体N_2O释放与反硝化微生物丰度和组成的关系

不同粒径团聚体的物理化学和生物学特性差异可能影响N2O的产生与释放,但目前有关团聚体N2O释放的微生物学机制少有研究。本研究从菜地土壤中分筛出粒径为1 mm、2~4 mm和4~8 mm的团聚体并开展培养试验,通过实时荧光定量PCR(Quantitative Polymerase Chain Reaction,q PCR)与末端限制性片段长度多态性分析(Terminal-Restricted Fragment Length Polymorphism,T-RFLP)技术的结合,研究了不同粒径团聚体反硝化微生物群落数量与组成的变化规律及其与N2O释放的相互关系。结果表明,不同粒径团聚体N2O释放速率表现为:粒径小于1 mm团聚体2~4 mm团聚体4~8 mm团聚体;硝酸还原酶基因(nar G)和氧化亚氮还原酶基因(nos Z)的丰度也表现为小粒径团聚体最高,随团聚体粒径增加而显著下降。然而,不同粒径团聚体间含nar G和nos Z基因的群落组成并没表现出显著差异。因此,不同粒径团聚体N2O释放速率差异与反硝化功能微生物丰度密切相关,而与它们的组成没有显著相关性。     

N2O and NO fluxes between a Norway spruce forest soil and atmosphere as affected by prolonged summer drought

Global change scenarios predict an increasing frequency and duration of summer drought periods in Central Europe especially for higher elevation areas. Our current knowledge about the effects of soil drought on nitrogen trace gas fluxes from temperate forest soils is scarce. In this study, the effects of experimentally induced drought on soil N₂O and NO emissions were investigated in a mature Norway spruce forest in the Fichtelgebirge (northeastern Bavaria, Germany) in two consecutive years. Drought was induced by roof constructions over a period of 46 days. The experiment was run in three replicates and three non-manipulated plots served as controls. Additionally to the N₂O and NO flux measurements in weekly to monthly intervals, soil gas samples from six different soil depths were analysed in time series for N₂O concentration as well as isotope abundances to investigate N₂O dynamics within the soil. N₂O fluxes from soil to the atmosphere at the experimental plots decreased gradually during the drought period from 0.2 to −0.0 μmol m⁻² h⁻¹, respectively, and mean cumulative N₂O emissions from the manipulated plots were reduced by 43% during experimental drought compared to the controls in 2007. N₂O concentration as well as isotope abundance analysis along the soil profiles revealed that a major part of the soil acted as a net sink for N₂O, even during drought. This N₂O sink, together with diminished N₂O production in the organic layers, resulted in successively decreased N₂O fluxes during drought, and may even turn this forest soil into a net sink of atmospheric N₂O as observed in the first year of the experiment. Enhanced N₂O fluxes observed after rewetting up to 0.1 μmol m⁻² h⁻¹ were not able to compensate for the preceding drought effect. During the experiment in 2006, with soil matric potentials in 20 cm depth down to −630 hPa, cumulative NO emissions from the throughfall exclusion plots were reduced by 69% compared to the controls, whereas cumulative NO emissions from the experimental plots in 2007, with minimum soil matric potentials of −210 hPa, were 180% of those of the controls. Following wetting, the soil of the throughfall exclusion plots showed significantly larger NO fluxes compared to the controls (up to 9 μmol m⁻² h⁻¹ versus 2 μmol m⁻² h⁻¹). These fluxes were responsible for 44% of the total emission of NO throughout the whole course of the experiment. NO emissions from this forest soil usually exceeded N₂O emissions by one order of magnitude or more except during wintertime.     

Plant canopy effects on litter accumulation and soil microbial biomass in two temperate forests

The objective of this study was to determine whether differences in canopy structure and litter composition affect soil characteristics and microbial activity in oak versus mixed fir-beech stands. Mean litter biomass was greater in mixed fir-beech stands (51.9t ha⁻¹) compared to oak stands (15.7t ha⁻¹). Canopy leaf area was also significantly larger in mixed stands (1.96m² m⁻²) than in oak stands (1.73m² m⁻²). Soil organic carbon (C _org) and moisture were greater in mixed fir-beech stands, probably as a result of increased cover. Soil microbial biomass carbon (C _mic), nitrogen (N _mic), and total soil nitrogen (N _tot) increased slightly in the mixed stand, although this difference was not significant. Overall, mixed stands showed a higher mean C _org/N _tot ratio (22.73) compared to oak stands (16.39), indicating relatively low rate of carbon mineralization. In addition, the percentage of organic C present as C _mic in the surface soil decreased from 3.17% in the oak stand to 2.26% in the mixed stand, suggesting that fir-beech litter may be less suitable as a microbial substrate than oak litter.     

Depth of root symbiont occurrence in soil

Summary The woody legume Prosopis glandulosa (mesquite) growing in the California Sonoran Desert develops functional root symbiotic associations (N₂-fixing nodules, vesicular-arbuscular mycorrhizal fungi) at depths greater than 4 m in moist soil above a seasonally stable water table. Population densities of symbiotic microorganisms are substantially greater at depth than near the surface. Inferences of plant symbiotic dependence based upon examination of surface roots and soil may be incorrect since deep roots can support the symbioses which are critical for plants utilizing deep water.