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Mycorrhizae and PLANT HEALTH Edited by F Mycorrhizae AND PLANT HEALTH Edited by F. L. Pfleger and R. G. Lindennan SYMPOSIUM SERIES ROLE OF VAM FUNGÍ IN BIOCONTROL Robert G. Linderman USDA-ARS Horticultural Crops Research Laboratory Corvallis, Oregon Most terrestrial plant species have in their roots a symbiotic association with soil fungí called mycorrhiza. There are several categories of mycorrhizae, of which the largest group, vesicular- arbuscular mycorrhizae (VAM), form with most agricultural crops. VAM fungi exist in soil as thick- wal-led chlamydospores, or as vegetativa propagules in roots, that germinate in the rhizosphere/rhizopiane. Their hyphae penétrate the root cortex, ramifying intercellularly from the point of penetration. The fungus forms special haustoria-1ike structures (arbuscules or coiled hyphae) within cortical cells, separated from the host cytoplasm by the host plasma membrane and the fungal cell wall. Arbuscules provide increased surface área for metabolic exchanges between the host and fungal partners. VAM fungi also develop extraradical hyphae that grow into the surrounding soil, increasing the potential of the root system for nutrient and water absorption, and contributing greatly to the improvement of soil structure for better aeration and water percolation. New survival spores are usually borne on the extraradical hyphae, although spores of some fungal species are produced primarily intraradically. Generally, VAM cause few changes in root morphology, but the physiology of the host plant changes significantly. For example, tissue concentrations of growth regulating compounds and other chemical constituents change, photosynthetic rates increase, and the partitioning of photosynthate to shoots and roots changes (16). The improved 2 CHAPTER ONE potential for mineral uptake from the soil accounts generally less severe on VAM p for changes in the nutritional status of the host plants, but the responses may tissues, in turn changing structural and biochemical involved are controversial (42 aspects of root cells. This can alter membrane nematode infection are general permeability and thus the quality and quantity of root but not always, nematode popul exudation. Altered exudation induces changes in the indicated by number of galls, composition of microorganisms in the rhizosphere soil, unit root 1ength)(41). Atilai now appropriately called the "mycorrhizosphere" showed an increase in nematodi (51,67,82). The net effect of these changes is a plants. These differences ma; healthier plant, better able to withstand differences between nematode environmental stresses (52) and tolérate or reduce the be due to differences in VAM effects of plant diseases. colonization. Furthermore, a The purposes of this review are (a) to discuss with fungal pathogens, timing the role of VAM in the expression of plant disease and :nportant. the mechanisms involved therein, and (b) to discuss A number of mechanisms of factors that influence the role of VAM in biological and nematode pathogens have b control of plant diseases. evidence supporting each is r proposed mechanisms depend or host physiology. Changes in VAM EFFECTS ON FUNGAL ROOT PATHOGENS plants may change the attrad Since VAM fungi are major components of the 'e^iatode pathogens. VAM may rhizosphere, it is logical that they could affect the and thus reduce yield losses incidence and severity of root diseases. Whether they infection, especially in low do or not has been the subject of numerous reviews established early in the groi over the last 15 years (18,32,46,71,72,73), but there infection. This mechanism o is still controversy. Much of the 1Herature suggests the reduced nematode respons that VAM fungi reduce soilborne disease or the effects lowever, by work that showed of disease caused by fungal pathogens. Dehne (32) (55). Cooper and Grandison reported that disease damage was reduced in 17 out of variable by using P-tolerant the 32 reports cited. However, the reports are still :er high P conditions. St mixed, with some indicating no effect of the VAM effects, leading these worke fungus on disease (6,7,28,84,85), and others increased resistance to nems indicating increased disease severity (29,30,31). to improved host nutrition, Drawing conclusions is difficult, partly because so other physiological changes many different pathogens and diseases have been Strobel et al. (78) and 01 \\g split root techniques involved, and partly because of the experimental conditions of each study. Clearly, one should expect -eT.atode infection on VAM p varied results, even if the VAM fungi used had been two were together on the sai the same (71). suggested that competition involved, a mechanism suppo and Sikora (69), Suresh et VAM EFFECTS ON PATHOGENIC ROOT NEMATODES ükora (75), and MacGuidwin Root infections by pathogenic nematodes are studies indicated that nema LINDERMAN 3 >take from the soil accounts generally less severe on VAM plants than on nonVAM lonal status of the host plants, but the responses may vary, and the mechanisms «9 structural and biochemical involved are controversia! (42,44). Symptoms of 5 can alter membrane nematode infection are generally reduced, and often, ity and quantity of root but not always, nematode populations are reduced (as ion induces changes in the indicated by number of galls, juveniles or eggs per sms in the rhizosphere soil unit root length)(41). Atilano et al. (3), however, mycorrhizosphere" showed an increase in nematode populations on VAM p of these changes is a plants. These differences may be due largely to >le to withstand differences between nematode pathogens, but could al so i and tolérate or reduce the be due to differences in VAM fungi and their levéis of colonization. Furthermore, as with VAM interactions review are (a) to discuss with fungal pathogens, timing of VAM formation is sion of plant disease and important. •erein, and (b) to discuss A number of mechanisms of interaction between VAM í of VAM in biológica! and nematode pathogens have been considered, and the evidence supporting each is reasonable. All the proposed mechanisms depend on VAM-mediated changes in UNGAL ROOT PATHOGENS host physiology. Changes in root exudation by VAM plants may change the attractiveness of roots to components of the nematode pathogens. VAM may improve host plant vigor, they could affect the and thus reduce yield losses caused by nematode seases. Whether they infection, especially in low P soils and if VAM are F numerous reviews established early in the growth cycle, before nematode 71,72,73), but there infection. This mechanism of enhanced P nutrition in :he literature suggests the reduced nematode response has been challenged, Hsease or the effects however, by work that showed no effect from adding P logens. Dehne (32) (55). Cooper and Grandison (24,25) eliminated P as a reduced in 17 out of variable by using P-tolerant VAM fungi on plants grown the reports are still under high P conditions. Still, VAM reduced nematode Ffect of the VAM effects, leading these workers to conclude that ')» and others increased resistance to nematodes was not entirely due íverity (29,30,31) to improved host nutrition, but must involve some - Partly because so other physiological changes in the roots. Studies by ises have been Strobel et al. (78) and Oliveira and Zambolim (61) :he experimental using split root techniques indicated that reduced •ly, one should expect nematode infection on VAM plants only occurred if the fungí used had been two were together on the same roots. These results suggested that competition for food or space was involved, a mechanism supported by results of Salen GENIC ROOT NEMATODES and Sikora (69), Suresh et al. (79), Sitaramaiah and Sikora (75), and MacGuidwin et al. (55). These ogenic nematodes are studies indicated that nematode size was reduced and 4 CHAPTERONE rate of development of infection was slower 1n VAM plant due to changes in the ho roots than nonVAM roots. Physiological changes in VAM 2) pointed out, foliage dise roots could also change resistance to nematodes by oblígate and non-obligate leaf increased production of ínhibitory substances (79), or increased on VAM compared to n by changes in root exudation which could alter to enhanced development of the mycorrhizosphere populations and affect nematode to increased incidence or freq populations and survival. >at effect was correlated wit While the mechanisms are still controversial, the igher physiological activitie evidence strongly indicates that VAM suppress nematode plants (32,73). infections of roots or reduce nematode effects on plant growth and yield. Undoubtedly, the effects and MECHANISMS OF VAM EFFEC1 the mechanisms involved depend on the conditions of the test, the host plant, edaphic conditions, and the Since VAM have such a sig? species of VAM fungus involved. Nonetheless, it seems plant physiology and on bioloc safe to say that VAM do play a role in the biological "izosphere soil, it follows ^ control of root nematode diseases. the incidence and severity of role played by VAM in the bio "seases has been the subject VAM EFFECTS ON BACTERIAL DISEASES 5.18,32,41,46,70,71,72,73), The effects of VAM on bacterial diseases have not "terpretations have preclude been explored to any great extent. However, Garcia- ;hat VAM always suppress plan Garrido and Ocampo (36) showed that VAM tomato -nconsistencies should be exp (Lycopersicon Miller) plants exhibited greater growth cansidering the diverse exper than nonVAM plants inoculated with Pseudomonas use of different VAM fungi on synngae pv. syringae van Hall
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