A poly(A) ribonuclease controls the cellotriose-based interaction between Piriformospora indica and its host Arabidopsis

Piriformospora indica, an endophytic root-colonizing fungus, efficiently promotes plant growth and induces resistance to abiotic stress and biotic diseases. The fungal cell wall extract induces cytoplasmic calcium [Ca2+]cyt elevation in host plant roots. Here, we show that an elici-tor-active cell wall moiety, released by P. indica into the medium, is cellotriose (CT). CT in-duces a mild defense-like response including the production of reactive oxygen species, changes in membrane potentials and the expression of genes involved in growth regulation and root development. CT based [Ca2+]cyt elevation in Arabidopsis roots does not require BAK1 coreceptor, or the putative Ca2+ channels TPC1, GLR3.3, -2.4 and -2.5 and operates synergistically with the elicitor chitin. We identified an ethylmethane-sulfonate-induced mu-tant ([Ca2+]cyt elevation mutant, cycam) impaired in response to CT, cellooligomers (n = 2, 4-7), but not to chitooligomers (n = 4-8) in roots. The mutant contains a single nucleotide ex-change in the gene encoding for a poly(A) ribonuclease (AtPARN, At1g55870) which de-grades poly(A) tails of specific mRNAs. The wild-type PARN cDNA, expressed under the control of a 35S promoter, complements the mutant phenotype. Our finding of cellotriose as a novel chemical mediator might help to understand the complex P. indica-plant mutual rela-tionship in beneficial symbiosis.