Tripartite associations in snowbrush (Ceanothus velutinus): effect of vesicular–arbuscular mycorrhizae on growth, nodulation, and nitrogen fixation

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Abstract
Symbiotic associations were established between nitrogen-fixing nonleguminous (actinorrhizal) snowbrush (Ceanothus velutinus Dougl.) seedlings and two categories of microorganisms: vesicular–arbuscular (VA) mycorrhizal fungi and a filamentous actinomycete capable of inducing nodule formation. The actinomycete is housed in nodules where fixation of atmospheric dinitrogen occurs and is made available to the host plant; the mycorrhizal fungus is both inter- and intra-cellular within the root tissue and may be found within the nodules. The two major nutrients, N and P, are made available and can be supplied to the host plant by these two symbiotic microorganisms. The root system of snowbrush seedlings was dually colonized by VA mycorrhizal fungi and a nitrogen-fixing actinomycete and the possibility of a direct interaction between the endophytes in the symbioses was investigated. Dually infected plants showed increases in total dry weight of shoots and roots, number of nodules, weight of nodular tissue, as well as higher levels of N, Ca2+, and P, and an increase in nitrogenase activity as measured by acetylene reduction.