Summary
Growth and activity at low temperatures and possible physiological and ecological mechanisms underlying survival of fungi isolated from the cold Arctic and Antarctic are reviewed here. Physiological mechanisms conferring cold tolerance in fungi are complex; they include increases in intracellular trehalose and polyol concentrations and unsaturated membrane lipids as well as secretion of antifreeze proteins and enzymes active at low temperatures. A combination of these mechanisms is necessary for the psychrotroph or psychrophile to function. Ecological mechanisms for survival might include cold avoidance; fungal spores may germinate annually in spring and summer, so avoiding the coldest months. Whether spores survive over winter or are dispersed from elsewhere is unknown. There are also few data on persistence of basidiomycete vs microfungal mycelia and on the relationship between low temperatures and the predominance of sterile mycelia in tundra soils. Acclimation of mycelia is a physiological adaptation to subzero temperatures; however, the extent to which this occurs in the natural environment is unclear. Melanin in dark septate hyphae, which predominate in polar soils, could protect hyphae from extreme temperatures and play a significant role in their persistence from year to year.

https://nph.onlinelibrary.wiley.com/doi/pdf/10.1046/j.1469-8137.2001.00177.x

Cold adaptation in Arctic and Antarctic fungi

This is not a review paper in the traditional sense, of which there are many. Three of the most influential reviews that summarized well some of the “older” literature include those by Nicolson (1967), Gerdemann (1968) and Mosse (1973). Instead, in this brief and incomplete work, we attempt to show the historical development of research on arbuscular mycorrhizas. We owe much to those who have written other historical accounts, including Rayner (1926–1927), Trappe and Berch (1985), Mosse (1985), Schenck (1985), Harley (1991) and Allen (1996), but the contents of this work naturally reflect our own ignorance, interests and biases. It was often difficult to distinguish between the historical and the contemporary, and we did not use any specific cutoff date in making this distinction. The degree to which we include “contemporary” literature was determined by our own assessment of its connectedness to older literature. In any case, we hope this will be of some interest to those of you who study the arbuscular mycorrhiza, and that it will serve the purpose of providing what we consider to be an important historical context for current researchers. We wish you good fortune in your research.

A history of research on arbuscular mycorrhiza

Arbuscular mycorrhizal (AM) fungi are ubiquitous in agricultural soils. These fungi play important roles in plant nutrition and soil conservation. The persistence of AM fungi in ecosystems depends on the formation and survival of propagules (e.g., spore, hyphae and colonized roots). While spores are considered to be resistant structure that may be view as “long-term” propagules when viable host plants are not present, hyphae are considered to be the main source of inocula when host plants are present and the soil is not disturbed. Tillage is an integral part of modern agriculture that can modify the physical, chemical and biological properties of a soil. Consequently, tillage practices may also affect AM fungi. The various tillage practices used in the management of soil for maximum crop production may negatively impact the survival of AM fungal propagules. In tilled soil, certain AM species may survive while others may disappear. Because AM fungi are more abundant in the topsoil, deep plowing may dilute ...

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Tillage or no-tillage: Impact on mycorrhizae

It is often assumed that all species of arbuscular mycorrhizal fungi (AMF) have the same function because of the ubiquity of the arbuscular mycorrhizal symbiosis and the fact that all AMF occupy the same plant/soil niche. Despite apparent differences in the timing of evolutionary divergence and the morphological characteristics of AMF from the different genera, the majority of studies on these fungi use only species of Glomus. There is increasing evidence, however, that the mechanisms involved in the establishment of a mycorrhiza may differ for species and genera of AMF and influence their subsequent function. The aim of this paper is to highlight the diversity in the form and function of AMF from different genera, knowledge of which is vital in understanding their ecological roles. Potential use of biochemical and molecular approaches to detect AMF in planta and ex planta is also discussed.

Mycelium of Arbuscular Mycorrhizal fungi (AMF) from different genera: form, function and detection

Microbial communities in plant roots provide critical links between above- and belowground processes in terrestrial ecosystems. Variation in root communities has been attributed to plant host effects and microbial host preferences, as well as to factors pertaining to soil conditions, microbial biogeography and the presence of viable microbial propagules. To address hypotheses regarding the influence of plant host and soil biogeography on root fungal and bacterial communities, we designed a trap-plant bioassay experiment. Replicate Populus, Quercus and Pinus plants were grown in three soils originating from alternate field sites. Fungal and bacterial community profiles in the root of each replicate were assessed through multiplex 454 amplicon sequencing of four loci (i.e., 16S, SSU, ITS, LSU rDNA). Soil origin had a larger effect on fungal community composition than did host species, but the opposite was true for bacterial communities. Populus hosted the highest diversity of rhizospheric fungi and bacteria. Root communities on Quercus and Pinus were more similar to each other than to Populus. Overall, fungal root symbionts appear to be more constrained by dispersal and biogeography than by host availability.

Plant host and soil origin influence fungal and bacterial assemblages in the roots of woody plants

The present investigation examines (1) whether the external VAM mycelium survives winter freezing to act as a source of inoculum in the spring, and (2) whether soil disturbance reduces the infectivity of the external VAM mycelium following freezing of the soil. Sealed pouches of fine nylon mesh were placed in pots containing soil inoculated with a Glomus species. The mesh was impervious to roots but not to hyphae. Following two 3-week growth cycles of maize in the pots, the pouches were transplanted to the field. Pouches were removed from the field once during the 4 months when the soil was frozen, and once after spring thaw. Measurements were made of VAM spore density, hyphal length and viability in the pouches. Bioassays for infectivity were conducted on all pouches. Some VAM hyphae survived freezing and remained infective following winter freezing, in the absence of plant roots. Soil disturbance did not reduce the infectivity of hyphae following exposure to freezing temperatures. We observed a change in the distribution of viable cytoplasm within hyphae over winter, which we hypothesize represents an adaptation allowing hyphae to survive freezing temperatures. We suggest that the effect of disturbance on hyphal infectivity may be related to this seasonal change in the distribution of hyphal viability.

Survival of the external mycelium of a VAM fungus in frozen soil over winter

SUMMARY
Abuscular mycorrhizal fungi are thought to survive adverse environmental conditions primarily as spores. Extraradical mycelia of two Glomus species were produced in fine mesh pouches which excluded roots but not hyphae. The mycelia in these pouches were exposed to freezing conditions, either in the field or in a controlled–temperature chamber. Bioassay plants were grown directly in the pouches and mycorrhizal colonization was assessed after 1 month. The mycelia remained infective in frozen soil over winter. This survival was not dependent on either the presence of root pieces or on the connection of mycelia to roots. Spores were not an effective inoculum in these bioassays. Overwinter survival of mycelia would enable plants to become incorporated into functional mycorrhizal associations early in spring.

H. D. Addy

Papers

Survival of the external mycelium of a VAM fungus in frozen soil over winter

Infectivity of the propagules associated with extraradical mycelia of two AM fungi following winter freezing