An analysis of the current state of knowledge of symbiotic fungal associations in 'lower' plants is provided. Three fungal phyla, the Zygomycota, Ascomycota and Basidiomycota, are involved in forming these associations, each producing a distinctive suite of structural features in well-defined groups of 'lower' plants. Among the 'lower' plants only mosses and Equisetum appear to lack one or other of these types of association. The salient features of the symbioses produced by each fungal group are described and the relationships between these associations and those formed by the same or related fungi in 'higher' plants are discussed. Particular consideration is given to the question of the extent to which root fungus associations in 'lower' plants are analogous to 'mycorrhizas' of 'higher' plants and the need for analysis of the functional attributes of these symbioses is stressed. Zygomycetous fungi colonize a wide range of extant lower land plants (hornworts, many hepatics, lycopods, Ophioglossales, Psilotales and Gleicheniaceae), where they often produce structures analogous to those seen in the vesicular-arbuscular (VA) mycorrhizas of higher plants, which are formed by members of the order Glomales. A preponderance of associations of this kind is in accordance with palaeohbotanical and molecular evidence indicating that glomalean fungi produced the archetypal symbioses with the first plants to emerge on to land. It is shown, probably for the first time, that glomalean fungi forming typical VA mycorrhiza with a higher plant (Plantago lanceolata) can colonize a thalloid liverwort (Pellia epiphylla), producing arbuscules and vesicles in the hepatic. The extent to which these associations, which are structurally analogous to mycorrhizas, have similar functions remains to be evaluated. Ascomycetous associations are found in a relatively small number of families of leafy liverworts. The structural features of the fungal colonization of rhizoids and underground axes of these plants are similar to those seen in mycorrhizal associations of ericaceous plants like Vaccinium. Cross inoculation experiments have confirmed that a typical mycorrhizal endophyte of ericaceous plants, Hymenoscyphus ericae, will form associations in liverworts which are structurally identical to those seen in nature. Again, the functional significance of these associations remains to be examined. Some members of the Jungermanniales and Metzgeriales form associations with basidiomycetous fungi. These produce intracellular coils of hyphae, which are similar to the pelotons seen in orchid mycorrhizas, which also involve basidiomycetes. The fungal associates of the autotrophic Aneura and of its heterotrophic relative Cryptothallus mirabilis have been isolated. In the latter case it has been shown that the fungal symbiont is an ectomycorrhizal associate of Betula, suggesting that the apparently obligate nature of the association between the hepatic and Betula in nature is based upon requirement for this particular heterotroph.

Examination of the roots of land plants has revealed the occurrence of mycorrhiza in the majority of species, over 70% of which are hosts to zygomycetous fungi that form vesicular–arbuscular (VA) associations. On the basis of experiments with a small number of host species showing enhancement of growth following colonization, it is widely assumed that wherever mycorrhizas are observed, the symbiosis is of the mutualistic type. The value of definitions based on structural rather than functional attributes is here brought into question by experiments simulating the ecologically realistic circumstance in which seeds germinate in soil in the presence or absence of established VA mycelium. These reveal a spectrum of fungal impacts in which some species respond mutualistically, while others, putative hosts or nonhosts, are antagonised, showing reduction of yield and survivorship and, hence, a loss of fitness relative to plants grown without VA fungi. Antagonised species normally grow in disturbed, open habitats and fail to establish in closed communities. It is hypothesised that their turf incompatibility arises from a sensitivity to interference by VA fungi, which consigns them to ruderal habitats. Mycorrhizal fungi, thus, play a role in defining the ecological niches occupied by plants and in determining of plant community composition. Key words: mycorrhiza, vesicular–arbuscular, mutualism, symbiosis, antagonism, plant community.

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While it is now widely accepted, even by ecologists, that most plants in the majority of ecosystems are infected by mycorrhizal fungi, few experiments have been designed to investigate the function of the mutualism at the community level. Those involved with mycorrhizal research have been largely preoccupied with questions of the mineral, particularly phosphorus, nutrition of individual plants, while plant community ecologists have too often found it convenient, even when acknowledging the presence of infection, to ignore its possible function in the ecosystem.This presentation examines a selected number of seminal papers written by plant community ecologists and highlights some of 'the most striking mysteries' which they reveal. It describes experiments designed to determine whether knowledge of the presence and activity of the mycorrhizal mycelium can help us to unravel the 'mysteries' which they define.It is revealed that by having direct adverse effects upon seedlings of many 'r' selected species, while at the same time being beneficial, if not essential, to those that are 'K' selected, the activities of the mycelium of VA fungi have a direct bearing upon community composition. The extent to which 'turf compatibility' is actually a reflection of the compatibility of plant species with the VA mycorrhizal mycelium is discussed and the possible role of the mycelium in consigning some species to the ruderal habit is considered.It is concluded that those attempting scientifically to understand, or managerially to manipulate, plant communities, without recognizing the role of the mycorrhizal mycelium, do so at their peril, and it is recommended that scientists involved in research on mycorrhiza extend their vision beyond the limited horizons which are currently so often defined by considerations of the phosphorus nutrition of individual host plants.

A major problem affecting grassland productivity in the semiarid southwestern USA is the low quantity of soil organic matter and plant-available N. In this study, dried, anaerobically digested sewage sludge was applied at three rates (22.5, 45, and 90 Mg ha~') to a degraded semiarid grassland site to determine the effects of sludge on soil chemical and heavy metal properties, and vegetative yields and quality over two growing seasons. Most nutrients, including soil N, P, and K, increased linearly as a result of sludge amendment. Soil pH decreased linearly from 7.8 to 7.4 with the application of sludge after the second growing season, but did not significantly increase the solubility of soil heavy metals (Pb and Cd). On the other hand, the levels of DTPA-extractable soil micronutrients (i.e., Cu, Mn, Zn) increased linearly with sludge rates to levels recommended for adequate plant growth. Total herbaceous plant yields increased significantly with sludge amendment, particularly at the 45 Mg ha' 1 sludge rate, as compared with the control treatment. Blue grama [Bouteloua gracilis (H.B.K.) Lag. ex Steud.] yields increased two-to threefold in amended plots over the unamended control after two growing seasons. The forage quality of blue grama, galleta [Hilaria jamesii (Torr.) Benth.], and bottlebrush squirreltail \Sitanion hystrix (Nutt.) J.G. Sm.], improved significantly as the levels of tissue N, P, and K, and crude protein increased linearly with sludge application. As expected, levels of Cd and Pb in all plant tissues did not increased significantly as a result of sludge amendment. All micronutrients increased to acceptable levels in deficient plant tissues. Sewage sludge applied at the rate of 45 Mg ha ' gave the most favorable soil and vegetation results. P.R. Fresquez, 2205 Columbia SE, Albuquerque, NM 87106; and R.E. Francis and G.L. Dennis, USDA Forest Service, Rocky Mountain Forest and Range Exp. St., Albuquerque, NM 87106.