Long-term suppression of Pythium ultimum in arid soil using fresh and composted municipal wastes

 The effect of addition of municipal solid waste (MSW) at different degrees of stabilisation on the biological properties of an arid soil was studied 24 months after application. This included effects on the indigenous soil microflora and soil enzyme activities in the presence and absence of Pythium ultimum. The addition of organic waste (fresh or composted) reduced populations of culturable bacteria and fungi and disease symptoms caused by P. ultimum, and resulted in heavier plants with longer and more extensively branched roots. Addition of organic waste increased the population size of culturable bacteria and fungi, while enzymatic activity of the soil was higher in soil amended with organic matter than in non-amended soil. Populations of biological control agents, such as Trichoderma and fluorescent pseudomonads, were larger in soil amended with organic matter. The addition of urban waste could therefore be a suitable technique with which to restore soil quality by stimulating biological control against plant pathogens such as P. ultimum. Received: 3 May 1999     

Surface-bound phosphatase activity in ectomycorrhizal fungi: a comparative study between a colorimetric and a microscope-based method

For the quantification of surface-bound phosphomonoesterase activity (SBPA) of fungi, roots, or mycorrhiza, a colorimetric method based on p-nitrophenyl phosphate (pNPP) is widely used. Unfortunately, this method does not reveal information about the localization of the surface-bound phosphomonoesterase (SBP). We introduce a method that localizes and quantifies SBPA in living hyphae of ectomycorrhizal fungi using confocal laser scanning microscopy of the hydrophilic substrate enzyme-labelled fluorescence (ELF-97) and compare it to the pNPP assay. ELF-97 turns into a strongly fluorescent precipitate upon activation by SBPA and forms bright fluorescent centres on the outer cell wall of the hyphae. Our data show that the enzymatic reaction is not substrate-limited during an incubation period of 15 min in fungal hyphae of Pisolithus tinctorius, Cenococcum geophilum, and Paxillus involutus. Image-processing routines determined the total intensity and the average number of the fluorescent ELF-97 centres per micrometre fungal hyphae of C. geophilum and Paxillus involutus. ELF-97 and pNPP detected similar variations of the SBPA at different pH values (3–7) during the measurement and different phosphorus (P) concentrations during the growth period of the fungi. The ELF-97 method revealed that C. geophilum and Paxillus involutus adapt in different ways to the variation of the P concentrations during the growth period by varying the number, the activity, or both properties of the SBP centres. The phosphatases show peak activities at different pH values, so the response of the fungal mycelium to varying P concentrations in soils is pH selective. In conclusion, ELF-97 is a promising substrate to reveal SBPA and adaptation strategies on a structural–physiological level.     

Inter- and intraspecific feeding selectivity of Folsomia candida (Willem) (Collembola,Isotomidae) on fungi

Summary Selective grazing of fungi by soil microarthropods may affect decomposition rates of litter materials and the structure of microarthropod and fungal communities. We developed laboratory methods to assay feeding selectivity and investigated the preferences of the collembolan Folsomia candida on three fungi: Acremonium sp., Paecilomyces varioti, and Penicillium citrinum. F. candida showed stronger preference for Acremonium sp. than for P. varioti and P. citrinum. Oviposition site selection followed the same pattern. Actively metabolizing hyphae of Acremonium sp. and P. varioti were preferred over senescent hyphae, while spores of P. citrinum were preferred over active hyphae. If microarthropod preference for active hyphae is extensive, microarthropod regulation of decomposition could be more important than their biomass indicates. Furthermore, as the P. citrinum results indicate, mechanisms of microbial dissemination may include selective grazing.Contribution in part to International Symposium on Faunal Influences on Soil Structure, Edmonton, Alberta, Canada (11–13 June 1984), organized by W. B. McGill; and the annual meeting of the Ecological Society of America, Minneapolis, Minnesota (17–21 June 1985)     

Adaptation of the enchytraeid toxicity test for use with natural soil types

Assessments of chemical toxicity to soil invertebrates have been traditionally performed in formulated Standard Artificial Soil, or in standard LUFA 2.2 natural soil. Physical and chemical properties of these standard soils often do not represent the diversity of properties of natural soils and can be inadequate surrogates of exposure conditions of soil biota in the field. We investigated performance of three species of the genus Enchytraeus (E. albidus, E. crypticus, E. luxuriosus) using soil types covering a wide range of European and North American soil properties. Results showed that adult survival and juvenile production by E. albidus and E. luxuriosus were inhibited in acidic soils with pH ≤ 5, while less acidic soils having properties within following ranges: 2.5–8.0% OM, 6–26% clay, and 4–80% sand, could sustain survival and reproduction at or near validity criteria levels of enchytraeid toxicity test ISO/16387. Performance of E. crypticus in North American soils having properties within following ranges: 1.2–42% OM, 1–29% clay, and 4.4–8.2 pH, met or exceeded all validity criteria. Data established in our studies can be used for amending the validity criteria of current ISO/16387 when used with E. crypticus and OECD soil.     

Reproductive performance of the generalist predator Hypoaspis aculeifer (Acari: Gamasida) when foraging on different invertebrate prey

In this study, we assessed the influence of prey quality and prey biomass during a standardized 3-week test on adult survival and reproductive output of the predatory mite Hypoaspis aculeifer when fed one of six different diets: springtails (Folsomia candida and Folsomia fimetaria), a storage mite (Caloglyphus cf. michaeli), an oligochaete (Enchytraeus crypticus), a nematode (Turbatrix silusiae), and a 1:1:1 mix of F. candida:F. fimetaria:E. crypticus. Our results revealed that a single prey species may be nutritionally sufficient for a 3-week period, as H. aculeifer performed equally well, or better, on a diet based on a 1:1:1 mix of F. candida:F. fimetaria:E. crypticus. However, when fed C. cf. michaeli, H. aculeifer had a poor reproductive output (<200 juveniles) and a reduced survival (60–70%). Thus, investigators should validate their choice of prey prior to testing H. aculeifer performance during toxicant exposure.     

Biological control of the take-all fungus, Gaeumannomyces graminis, by Phialophora radicicola and similar fungi

Phialophora radicicola is an avirulent fungal root-parasite of grasses and cereals, with runner hyphae like those of Gaeumannomyces graminis. Weakly and non-pathogenic varieties of these fungi control the pathogens, G. graminis vars. tritici and avenue. Biology of these fungi is considered and the evidence for biological control and possible mechanisms reviewed; control is probably widespread in natural plant communities, and host-mediated, perhaps by induction of plant resistance mechanisms.Prospects for application of biological control seem best for P. radicicola var. graminicola established on grass crops, as this is already exploited in British agriculture. New evidence is presented on the effects of grassland factors on this fungus, especially sward composition, age, mineral nutrition and management practices: its population might often be limited by the rate of new root production to replace those with cortices already colonized. Prospects for control by seed inoculation with P. radicicola var. radicicola and G. graminis var. graminis also seem good, but possible dangers of introducing them into cereal cropping are emphasized. The weak pathogens might be used also for indirect control by establishing hyper-parasites or inducing disease suppression (like take-all decline) in soils, but there is no evidence for ‘Phialophora decline’, at least in well-managed grasslands. Finally, different biocontrols of take-all might be combined, and biological with chemical ones for ophiobolus patch disease of turf.P. radicicola var. graminicola has a slight beneficial effect on grass yield, even when the pathogens, G. graminis vars. tritici and arenae arc absent; this probably contributes to its abundance in natural grasslands in Britain. The scale of biological control by this and similar fungi might explain why, in their absence, effective plant resistance to G. graminis is uncommon in the Gramineae.     

Solubilization of Insoluble Inorganic Phosphate by Hyphal Exudates of Arbuscular Mycorrhizal Fungi

Increased phosphate (P) uptake in plants by arbuscular mycorrhizal (AM) fungi is thought to depend mainly on the extension of external hyphae into soil. On the other hand, it is known that the hyphae of some kinds of ectomycorrhizal fungi release organic acids into soil and that they dissolve the insoluble inorganic P. This study collected hyphal exudates of AM fungi within compartmentalized pot culture and clarified their ability to solubilize insoluble inorganic P. Sterilized Andisol was packed in pots that were separated into root and hyphal compartments with a nylon net of 30 μm pore size. Seedlings of Allium cepa inoculated with AM fungi, Gigaspora margarita, or Glomus etunicatum were grown. Control pots were not inoculated. Mullite ceramic tubes were buried in the soil of each compartment and soil solution was collected. The anionic fraction of the soil solution was incubated with iron phosphate (4 mg FePO₄ in 1 mL of 0.4 acetate buffer). Solubilized P was measured. The AM colonization of plants inoculated with G. margarita and G. etunicatum was 86% and 54%, respectively. Adhesion of external hyphae was observed on the surface of the mullite ceramic tubes buried in soil of the hyphal compartment. Colonization of both fungi increased shoot P uptake and growth. Soil solution collected from the hyphal compartments of both fungi solubilized more P than did that from uninoculated plants. It is suggested that hyphal exudates can contribute to increased P uptake of colonized plants.     

Glomalin-related soil protein and water relations in mycorrhizal citrus (Citrus tangerina) during soil water deficit

Glomalin-related soil protein (GRSP), a glycoprotein of arbuscular mycorrhizal fungi (AMF) secreted into soil, governs the aggregate stability, but the role of GRSP in soil and plant water is sparsely studied. The 24-week-old red tangerine (Citrus tangerina) inoculated with Glomus etunicatum and G. mosseae were subjected to a soil drying for 12 days as soil water deficit (SWD). Length of SWD significantly reduced mycorrhizal colonization, soil hyphal length, and leaf and soil water potential (Ψ), but increased total GRSP (T-GRSP), easily extractable GRSP (EE-GRSP), and proportion of water-stable aggregates (WSAs) in >0.25 mm size, irrespective of AMF source. The AMF-inoculated seedlings showed significantly higher T-GRSP, EE-GRSP, and leaf/soil Ψ than non-AMF seedlings during SWD. A significantly positive correlation was observed for mycorrhizal colonization versus leaf or soil Ψ, and hyphal length versus leaf Ψ, suggesting that root intra- and extra-radical hyphae participated in water transport. Interestingly, in GRSP fractions, only T-GRSP was significantly positively correlated with 0.25–1 and >0.25 mm WSA and negatively with leaf and soil Ψ. These results revealed a strong glue function of T-GRSP (not EE-GRSP and hyphae) to alter the proportional distribution of WSA size, thereby aiding toward prevention of soil water loss for improving soil–plant water relations.     

Palatability of microfungi to soil arthropods in relation to the functioning of arbuscular mycorrhizae

We investigated the feeding preferences of six species of mites and collembolans for three fungi commonly associated with roots of Acer saccharum (Glomus macrocarpum, Alternaria alternata and Trichoderma harzianum), from a maple-forest soil in southern Ontario, Canada. Experiments were also conducted in vitro to determine animal feeding responses to (1) increasing quantities of hyphal biomass, (2) the presence of root vs. litter fungal substrates, and (3) hyphae of different widths of Glomus macrocarpum. The results indicate that arthropods prefer to graze in the litter region rather than in the deeper soil layers. Under ideal moisture/temperature conditions, animals are forced to the lower regions by interspecific interactions. They prefer to graze on hyphae of conidial fungi rather than on those of arbuscular mycorrhizal fungi. When arbuscular mycorrhizal fungal hyphae are grazed, there is a clear preference for the narrower hyphae, which are those further away from the root. The thicker hyphal segments, commonly found connecting absorptive hyphal fans to roots, were less preferred. These data are not consistent with the hypothesis that microarthropods are detrimental to arbuscular mycorrhizal associations, and suggest that Glomalean fungi may have evolved mechanisms to deter grazing by microarthropods.     

Differential production of oxalate by mycorrhizal fungi in arid ecosystems

Oxalate crystals and elements binding to the surfaces of mycorrhizal fungal hyphae were examined using scanning electron microscopy coupled with X-ray analysis of elemental composition. Mycorrhizae from the arid zone vegetation types in southern California were examined including chaparral, riparian oak woodlands, coastal sage, grasslands, and deserts. Only mat-forming ectomycorrhizal hyphae, such as Hysterangium separabile, were found to produce oxalate crystals. None of the arbuscular mycorrhizal fungal hyphae (Glomus spp. and Acaulospora elegans) examined had crystal structures associated with them. The hyphae of Hysterangium separabile without crystals did not show the Ca peaks that were present when the crystals existed nor did the arbuscular mycorrhizal fungal hyphae have the Ca peaks. The elimination of arbuscular mycorrhizae using benomyl did not affect soil P or oxalate. These data indicate that there are some fundamental differences in chemical exudation between mycorrhizal fungi that could affect P uptake and cycling in arid ecosystems. Received: 7 December 1994     

Interaction of bacteria-feeding soil flagellates and Pseudomonas spp.

Pseudomonas strains may be used as alternatives to fungicides as some of them produce secondary metabolites, which can inhibit growth of plant pathogenic fungi. Increased knowledge of non-target effects of the antagonistic bacteria on other soil organisms as well as of the survival and predation resistance of the antagonistic bacteria is necessary for risk assessment and increased performance of antagonistic bacteria as biological control agents. In the present study, we aimed to investigate the difference between Pseudomonas spp. with respect to their predation resistance to and effects on the three different and common soil flagellates Bodo caudatus, Cercomonas longicauda, and Neocercomonas jutlandica. Two antagonistic Pseudomonas: Pseudomonas fluorescens CHA0 and P. fluorescens DR54 and two positive control strains: P. fluorescens DSM 50090^T and Pseudomonas chlororaphis ATCC 43928 were studied. CHA0, which is reported to produce a range of different antibiotic substances, was inhibitory to all the tested flagellates whereas DR54, which produces viscosinamide, only inhibited the growth rate of B. caudatus. Removal of spent medium from a CHA0 culture reduced the growth inhibition of C. longicauda, although the growth rate was still lowered compared with DSM 50090^T. In contrast, removal of spent medium from a DR54 culture had minor effect on C. longicauda. The flagellate B. caudatus was far more sensitive towards the antagonistic Pseudomonas strains of the tested flagellates.     

Variable impacts of enchytraeid worms and ectomycorrhizal fungi on plant growth in raw humus soil treated with wood ash

An increasing amount of evidence shows the context dependent nature of various biotic interactions across terrestrial and aquatic ecosystems. We established a laboratory experiment to study whether the effects of Cognettia sphagnetorum (Enchytraeidae) and ectomycorrhizal fungi on Scots pine (Pinus sylvestris) seedling growth are influenced by wood ash application. Acidic coniferous forest soil was treated with wood ash at 5000 kg ha⁻¹ or left as ash-free control and inoculated with soil saprotrophic microbes and nematodes. The microcosms were destructively sampled 26 and 51 weeks after initiation of the experiment. We measured enchytraeid and pine seedling biomass, abundance of nematodes and leaching of NH₄⁺-N and NO₃⁻-N at both samplings, and root length and N concentration of pine needles at the end of the experiment. On average enchytraeids and mycorrhizal fungi enhanced pine biomass production in the ash-free control soils, however, their impact was most pronounced when these organisms were alone in the systems. In fact, mycorrhizas tended to have a negative impact on the seedlings in the presence of enchytraeids. Wood ash had a clear negative impact on enchytraeid populations. Wood ash decreased pine growth when enchytraeids and EM-fungi were alone in the systems, but when together they apparently compensated for the negative effects of wood ash on the seedlings. It is concluded that interactions between soil fauna, mycorrhizal fungi and plants are context dependent, thereby rendering predictions of the outcome of species interactions in soil food webs a demanding task.     

VA菌根对土壤酸度的耐性

A 45－day greenhouse experiment was carried out to determine effect of vesicular-arbuscular（VA）my-corrhizal fungi on colonization rate,plant height,plant growth,hyphae lenth,total Al in the plants,ex-changeable Al in the soil and soil pH by comparison at soil pH 3.5,4.5 and 6.0 Plant mung bean(Phaseolus radiatus L.)and crotalaria(Crotalaria muronata Desv.) were grown with and without VA mycorrhizal fungi in pots with red soil,Ten VA mycorrhizal fungi strains were tested,including Glomus epigaeum(No.90001),Glomus caledonium(No.90036),Glomus mosseae(No.90107),Acaulospora soo.(No.34),Scutellopora het-erogama(No.36),Scutellospora calopsora(No.37),Glomus manihotis(No.38).Gigaspora spp.(No.47),Glomus manihotis(No.49),and Acaulospora spp.(No.53).Being the most tolerant to acidity,strain 34 and strain 38 showed quicker and higher-rated colnization without lagging,three to four times more in number of nodules,two to four times more in plant dry weighy,30% to 60% more in hyphae length,lower soil exchaneable Al,and higher soil pH than without VA mycorrhizal fungi(CK).Other strains also could improve plant growth and enhance plant tolerance to acidity,but their effects were not marked.This indicated that VA mycorrhizal fungi differed in the tolerance to soil acidity and so did their inoculation effects.In the experiment,acidic soil could be remedied by inoculation of promising VA mycorrhizal fungi tolerant of acidity.     

丛枝菌根根外菌丝对铵态氮和硝态氮吸收能力的比较

采用空气隔板分室法并结合15N标记技术,以玉米为宿主植物并接种Glomus mosseae和Glomus intraradices,比较了这两种真菌根外菌丝对铵态氮和硝态氮吸收传递能力的差异。结果表明,丛枝菌根根外菌丝吸收传递氮的能力因菌种和氮素形态而异。两种真菌根外菌丝吸收传递NH4+-N能力均高于NO3--N;G. intraradices根外菌丝吸收传递氮的能力高于G. mosseae,这可能与两种真菌根外菌丝生长量有关。     

Mycorrhizal function on soil aggregate stability in root zone and root-free hyphae zone of trifoliate orange

Arbuscular mycorrhizal fungi (AMF) influence soil aggregate stability through their hyphae, roots, and glomalin-related soil protein (GRSP); however, the individual effect of these factors is difficult to distinguish. Pots separated by a 37-μm mesh bag buried in the middle of each pot was used to establish root zone (root + hyphae) and hyphae zone (roots free), where the Poncirus trifoliata seedlings were colonized by Funneliformis mosseae or Paraglomus occultum in root zone. AMF inoculation significantly increased shoot, root, and plant’s total biomass, soil organic carbon, GRSP fractions, 2–4 and 1–2 mm size water-stable aggregates, and mean weight diameter (MWD) in root or hyphae zone. Within root zone, root colonization and biomass presented stronger relationship with MWD than GRSP fractions. While, within hyphae zone, total of GRSP fraction was significantly correlated with MWD. The study, suggested further that root biomass and colonization were the main mechanisms in root zone for improving aggregate stability, whereas total of GRSP fractions was of paramount importance in hyphae zone. Mycorrhizal effect on aggregate stability was observed to be contrastingly different between root zone and hyphae zone.     

丛枝菌根真菌对西藏高原固沙植物吸磷效率的影响

采用盆栽方法,就外源菌种、土著菌种(含混合菌种)对固沙植物白草(Pennisetum.flaccidum)生长和吸磷效率的影响进行了研究。结果表明,白草具有较高的菌根依赖性(平均达166.4%);不同AM真菌(或真菌组合)对白草根系均具显著的侵染效应。随菌根侵染率的提高,植株生物量、吸磷量均呈显著增加(相关系数分别为0.7465*、0.6000*);菌根菌丝对白草吸收土壤磷素的贡献十分明显,各接种处理菌根菌丝对植物吸收土壤磷素的贡献量、贡献率分别在3.2～11.6.mg/pot和61.5%～85.3%之间;接种菌根处理植株吸磷量呈Glomus.intraradicesG.mosseae+G.etunicatum+G.intraradices+Scutellospora.erythropaG.mosseae(外源菌种)G.mosseae+G.intraradices+Scutellospora.calosporaG.mosseae-I(土著菌种)G.etunicatum的趋势。此外,不同AM真菌对寄主植物地上部、根部生物量和吸磷量的影响程度明显不同,一般呈地上部根系的趋势,但寄主植物根系的生长速率相对较快;土著菌种中,多菌混合接种对寄主植物的侵染效应明显高于单一接种。     

不同氮磷比例营养液对AM真菌生长发育的影响

温室条件下,研究不同氮磷比例营养液对丛枝菌根(Arbuscular.mycorrhizae,AM)真菌(Glomus.mosseae)生长发育的影响。结果表明:在一定磷水平条件下增加氮的水平,有利于菌根真菌的生长发育,但超过一定水平则会抑制菌根真菌的生长发育,说明氮水平也会影响菌根真菌的生长发育,这种影响与氮磷比例有关。综合比较菌根长度、根外菌丝量及孢子数三项指标,在20%浓度Hoagland营养液的基础上,将氮磷比例提高到4∶2有利真菌生长。在AM菌剂生产中,通过营养液中氮磷比例的调控能获得较大数量的侵染根段、菌丝及孢子等繁殖体。     

Dynamics and stratification of Enchytraeidae in the organic layer of a Scots pine forest

In a Dutch Scots pine forest an experiment was conducted to quantify the role of soil biota in the functioning of the soil ecosystem, and the effects of enhanced nitrogen deposition. For this, the site was sampled at 8-week intervals during 2.5 years. This paper reports on the population dynamics of enchytraeids in the field and in stratified litterbags. Mean yearly abundance of the enchytraeid community in the field was 47 600 m^–2, or 0.70 g (dry weight) m^–2. The community consisted mainly of three species: Cognettia sphagnetorum, Marionina clavata and Achaeta eiseni, of which C. sphagnetorum was dominant. The enchytraeid populations showed a marked stratification in the same sequence. Freshly fallen pine needles were colonized by C. sphagnetorum, while other species followed much later. It was found that data from the litterbags were reasonably comparable with field data, when expressed per gram of dry substrate, but less so when expressed per square metre. Multiple regression analysis of the data showed that the population dynamics in the litter layer could largely be explained by temperature and moisture fluctuations; in deeper layers other factors, such as the stage of decomposition, were probably more important. Received: 26 June 1997     

Earthworms and enchytraeids in conventional and no-tillage agroecosystems: A biocide approach to assess their role in organic matter breakdown

Summary Earthworm and enchytraeid densities and biomass were sampled over an 18-month period in conventional and no-tillage agroecosystems. Overall, earthworm densities and biomass in the no-till system were 70% greater than under conventional tilling, and enchytraeid densities and biomass in the no-till system were 50%–60% greater. To assess the role of annelids in the breakdown of soil organic matter, carbofuran was applied to field enclosures and target (earthworm and enchytraeid biomass, standing stocks of organic matter) and non-target effects (bacteria, fungi, protozoa, nematode and microarthropod densities, litter decay rates, plant biomass) were determined in two 10-month studies. In the winter-fall study, carbofuran reduced the annelid biomass, and total soil organic matter standing stocks were 47% greater under no-till with carbofuran compared to control enclosures. Twelve percent of the difference could have been due to non-target effects of carbofuran, as determined from litterbag decay rates. In the summer-spring study, carbofuran again significantly reduced the annelid biomass, and treated pens in the no-till area had significantly greater standing stocks of fine organic matter (43%–45%). Although the densities of bacteria and nematodes were reduced in carbofuran-treated litterbags under a no-till system, the rates of decay were not reduced and estimates of the amount of organic matter processed could not be adjusted for non-target effects. A 76% difference in the standing stock of coarse organic matter between control and carbofuran-treated pens in the conventional-till system indicated further non-target effects. We concluded that our estimates of the amount of organic matter processed by annelids in no-till and conventionally tilled agroecosystems represented a maximum potential because of the confounding non-target effects of carbofuran.     

Crop residue and Collembola interact to determine the growth of mycorrhizal pea plants

The effects of collembolan grazing on arbuscular mycorrhizal (AM) fungi and plant growth were studied in a controlled experiment utilizing a mix of AM fungi and the dominant collembolan species (Isotoma sp.) indigenous to the experimental soil. Collembolan (+/– Col) effects were examined in the presence and absence of crop residue (+/– Litter) incorporated into the experimental soil. Significant interactions between collembolans and crop residue occurred for mycorrhizal colonization of roots and plant growth. In the absence of crop residue, collembolans reduced root length colonized by AM fungi, total plant dry mass and seed pod yield. However, in the presence of crop residue, collembolans had no effect on root colonization by AM fungi, and increased total plant mass and pod yield. Crop residue increased root colonization by AM fungi, numbers of bacteria and saprophytic fungi (colony forming units), small- (<5 m) and large- (>5 m) diameter hyphal lengths in soil, and the final population of collembolans in soil. Collembolans reduced both small- and large-diameter hyphae in soil and the number of saprophytic fungi (colony forming units, p =0.052). Feeding preference experiments conducted in vitro showed that Isotoma sp. preferred to graze on mycorrhizal roots over nonmycorrhizal roots when given no other food choice. However, when crop residue was added as a food choice, Isotoma sp. showed a clear feeding preference for crop residue. We conclude that collembolan grazing on mycorrhizae can be detrimental to plant growth when other fungal food sources are limited, but grazing on mycorrhizal fungi does not occur when ample organic matter and associated saprophytic fungi are present in soils.