Tansley Review No. 76 Helper bacteria: a new dimension to the mycorrhizal symbiosis.

The symbiotic establishment of mycorrhizal fungi on plant roots is affected in various ways by the other microorganisms of the rhizosphere, and more especially by bacteria. This review discusses the case of some of these bacteria which consistently promote mycorrhizal development, leading to the concept of 'mycorrhization' helper bacteria (MHBs). Examples of MHB evidence are given from the literature, with special reference to the Douglas fir (Pseudotsuga menzeisii Mirb. Franco) Laccaria laccata Scop, ex Fr. ectomycorrhizal combination which has been more extensively studied. The fungal specificity of some MHBs and the various mechanisms underlying their effect are discussed, considering five hypotheses: effects on the receptivity of the root, effects on the root-fungus recognition, effects on the fungal growth, modification of the rhizospheric soil, and effects on the germination of the fungal propagule. MHBs are then considered for their ecological and evolutionary implications, and examples of practical applications in forest nurseries are given: when added to the fungal inoculum, MHBs can improve the success of ectomycorrhizal inoculation of planting stocks with fungi selected for their outstanding growth stimulation after outplanting. The conclusion points out a number of fundamental questions which remain unanswered about mycorrhization helper bacteria and suggests some investigation priorities in this new field of mycorrhiza research. Contents Summary 197 I. Introduction 197 II. Evidence for helper bacteria 198 III. Fungus-specificity of MHBs 200 IV. Mechanisms underlying the MHB effect 201 V. Ecological and evolutionary implications of MHBs 205 VI. Practical applications of MHBs 206 VII. Conclusions and perspectives 208 Acknowledgements 208 References 208.