FUNGISTASIS IN SOILS

SUMMARY 1. Fungistasis in soil is a widespread phenomenon affecting most fungal propagules, though some are insensitive. In most instances, it is coexistent with the presence of living microorganisms, and is annulled by energy-yielding nutrients. Fungistasis with characteristics similar to that in soil may also occur on leaves of plants. 2. Germination and growth of bacteria and actinomycetes is also restricted in soils. The characteristics of their inhibition appear to be the same as those for fungi. Therefore, the concept of a widespread microbial inhibition in soil can be applied to all three groups of microorganisms. 3. Fungistasis can be detected by various direct methods, or indirectly by methods involving the use of porous or permeable carriers. It may be expressed as a restriction on the final amount of germination (the usual parameter), germination rate (with time), and rate of germ-tube or hyphal growth. Since the expression of fungistasis is often complete in soil, titration with nutrients may be required to distinguish between the sensitivities of different fungi. 4. Fungistasis generally is expressed most strongly at soil moisture contents somewhat less than saturation. Its expression usually is maximal in neutral or slightly alkaline soils. In acidic conditions fungistasis may be lessened because of suppression of bacterial and actinomycete activity. Increased sensitivity of some fungi in soils of pH > 7.0 may be caused by a directly unfavourable effect of pH on the fungus. 5. Fungal species with small spores tend to be highly sensitive to fungistasis. These spores tend to germinate slowly and to require exogenous nutrients for germination. By contrast, species with larger spores and sclerotia often do not require exogenous nutrients for germination. The larger spores tend to germinate rapidly and to exhibit low sensitivity, as compared with small spores. A few nutrient-independent spores are insensitive to fungistasis. At least a part of the difference in sensitivity is related to germination time; spores which germinate slowly compete poorly with the soil micro-flora for their nutrients. 6. Fungistasis is often temporarily annulled by enriching the soil with energy-yielding nutrients. Usually, complex materials such as plant residues are most effective. A few weeks after such treatment, the level of fungistasis may, however, be increased. Annulment of fungistasis by compounds not utilized as energy sources has not yet been demonstrated. 7. Several soils naturally suppressive to Fusarium wilt diseases were more fungistatic to Fusarium than soils conducive to wilt. Potential means by which fungistasis may be manipulated to control root-infecting fungi are (a) through stimulation of germination with nutrients, thus exposing the germ tube to lysis, and (b) by increasing the fungistatic level of soil through appropriate amendments. 8. Volatile substances identified in soils, some of which are potentially inhibitory to fungi include (a) ammonia, which apparently is evolved from ammonium salts in some arid soils of high pH, (b) ethylene, which has been identified in some soils of pH < 7.0 (though high levels of this gas seem to be tolerated by most fungi), (c) allyl alcohol, and (d) other unidentified substances. Non-volatile inhibitors include high molecular weight substances revealed by molecular sieve chromatography of soil extracts. Microbial metabolites such as those present in staled fungal cultures also have been proposed to account for fungistasis. In a few soils fungistasis persists after sterilization because of the presence of inhibitory concentrations of calcium carbonate, iron or aluminium. Inherent in the proposition that inhibitory substances provide the primary mechanism of fungistasis is the concept of a highly complex phenomenon, involving various highly specific inhibitory and counteracting stimulatory substances, with the outcome for the fungus depending on the kinds and relative amounts of each present. 9. By the nutrient-deficiency hypothesis, the level of available nutrients in soil is insufficient to support germination of nutrient-dependent propagules, except in nutrient-rich microsites. Inhibition of nutrient-independent propagules is explained by loss of endogenous nutrients required for germination, through microbial nutrient competition. Evidence for this hypothesis is (a) the imposition of fungistasis on numerous nutrient-independent propagules during incubation on leaching model systems designed to simulate microbial nutrient competition in soil, (b) similar losses of endogenous nutrients occurring on soil and the leaching system, and (c) the fact that soils are chronically deficient in energy in relation to the microbial populations present, with the consequence that enforced inactivity is imposed upon most of the population at any given time for this reason alone, regardless of the presence or absence of fungistatic substances. Journal series article no. 7747 from the Michigan Agricultural Experiment Station.