Dimerization Controls the Activity of Fungal Elicitors That Trigger Systemic Resistance in Plants

Abstract

The soilborne fungus Trichoderma virens secretes a small protein (Sm1) that induces local and systemic defenses in plants. This protein belongs to the ceratoplatanin protein family and is mainly present as a monomer in culture filtrates. However, Hypocrea atroviride (the telomorph form of Trichoderma atroviride) secretes an Sm1-homologous protein, of the in and the in the of systemic in that and mainly as monomer and a in the of The that the to defenses in the form of and and the that the form of induces defenses in plants. that is as a the is and is as a in is a the of the and to a in the ceratoplatanin protein family to the of the and the of the to The that the the and the of systemic in plants. The soilborne fungus Trichoderma virens secretes a small protein (Sm1) that induces local and systemic defenses in plants. This protein belongs to the ceratoplatanin protein family and is mainly present as a monomer in culture filtrates. However, Hypocrea atroviride (the telomorph form of Trichoderma atroviride) secretes an Sm1-homologous protein, of the in and the in the of systemic in that and mainly as monomer and a in the of The that the to defenses in the form of and and the that the form of induces defenses in plants. that is as a the is and is as a in is a the of the and to a in the ceratoplatanin protein family to the of the and the of the to The that the the and the of systemic in plants. in a of in and to the a is and the in an the of as and and in the as and to as of is the of defenses the of systemic systemic culture systemic culture systemic in of the and systemic and However, the that the and the that This of to the of and to of the fungus Trichoderma The of virens virens to and defenses in The virens is the small protein to in the of plants. This is to and in in that of is in and the the that to the of the and and to of However, the to and the and the in to is a protein, a of to and belongs to the protein family mainly of and and is and of a of and virens culture as a protein However, in a to the form in The of monomer and of of in is the Hypocrea in and in culture and the the and the of the of and that the of the the of and to to a of and that the form of is as a However, present in of The of in and of the The of to that the the protein and the of the and the of defenses in plants. the of the of in the and in the of in plants. the protein to the to and the of is and Trichoderma in Trichoderma virens virens virens virens Trichoderma atroviride Trichoderma and Trichoderma the and as a in The in in a in a and in a a and of the to the of virens atroviride as The of the Trichoderma the of and and the and a and a a and to and of to and and of and virens and atroviride in culture of in in in the of The of and in the of and the and in the of and as in a the protein to the as a The as and and a of and and virens and atroviride in and a in and the protein a in the The of and and protein The the protein of and The protein in and the The of and and the form of in the the form of The and of the of and an The the and to an to the to the form of and the in The and in and to of and to The and The in a the a to the of and of and to a a of in and a in the protein the as the and the to the a and in the of in the of the and to the and a and The to to in virens and atroviride as a the of and as the The the The in a and of The in a the the The of in The and in The of the the (the of the of in a of of the and a and a of in a of in virens virens and atroviride a and and of and in a and in a a and the the in and as The of the the the of the The and of in in of form and an of monomer of and of and the of The in and the as the of the the in and a as in the the the and the and of the The of the the virens and atroviride in the of and to in the of of and and in the of plants. in the of of of atroviride of the in the of The of the of culture in the of in virens and atroviride in culture and in the of the in the of and the the of The of and in atroviride in the of However, in the atroviride in the of of and the in in the of is mainly a of a form and in virens and atroviride that the of and in virens the of virens and atroviride the of and the of in in the and of of and in and in the of in virens and and of in virens of the in The of of the of virens The the in of virens and atroviride to the systemic of the a is in a virens in the of of the in The to and in the The and the the and and However, virens the and a in the of the and and the of and The of the is in a virens and of in and of plants. Trichoderma as of The of and the in as the and the to The the of to a of virens atroviride virens atroviride in plants. The of the of and that virens virens atroviride the and of The the of the the The the in in to a of of the an to and to The the and of and virens and atroviride in the in and of and in a an of and an the the monomer of to the monomer of and the of and of the of an The the of and and the This the of to as that to a as the monomer and to The and in the of the and the protein in as The of and The that the of the in the of the and of and in the of a of and and and of However, in of and of and and of virens atroviride in and of and and of and The protein protein and the of The in and the of of the and to and the of the of the monomer and in and the of and and protein to monomer form of form of form of form of form of of and of and the of and the of and The in a that in The is in the is the the of in the and and and the form of in as a and the that the form of a of a the the of in of the and and and of of and protein in The is the and the to is The monomer and of in a and and and The in and of The the monomer form of form of form of of the in in a the of the in of the and a virens and to of the that a to of virens and in a a Trichoderma the of in the of of Trichoderma the of virens and as a a to that the the in of and the to the monomer and the and in the of the of to the monomer and a and the of a in and a to a monomer that the form of atroviride an in the The of is in the and of in the Trichoderma This is in Trichoderma and virens a to the to This is in and the is in of the protein of the of and in of to the is in and The a and a of of in Trichoderma and of in in of and to and virens virens virens atroviride of the of Trichoderma in a of and of and in that induces the present as a and to the that and in the of the of and the of the of the monomer and the and form of that the of and the monomer of that the form of protein the of the the of the The of the an of the as a the that the of an of the to that of the the that the form of a The and and The of the is in of the of and in and The of of monomer as a of and of as The to the The the of to a of of and the of the fungus and a of The the of the the The the in in to a of an to and to The the is a that in that protein is a in the of as and that the of an protein (Sm1) in in the of the that to the of in as the of in the form of the The of a of to and to the of of the protein the to defenses is the The of and that the of small to the in and the and virens and atroviride systemic in to in the of the of virens of the form of The of the form of atroviride as to However, that Trichoderma that in the that atroviride to the of form of in the of and as a in the However, virens in as as in and the of the monomer that of the the of the to that as of the the the of and to of the of as and as an in the of the of of to to the to a to and the of and to in the the that the to the a to and that the in is in plants. the of a the of the This the to and of the form that the of is to an the of Trichoderma and to and However, the in and that and of to and an protein in the and the of This the of a to to form a that the and a the and of the form of the of that the the to form in the of a of the to that This in virens and in the in is in an is This of in the an of the the of the in the However, in the of virens the of a the that in a protein in the of the as a monomer in the the and The form of the in to and to However, the is in the of the the to and in form the in the of in to in the in protein The present of a virens and an and to of the Trichoderma of and that is to virens This a virens and and the of the of the in of The and of in fungus and the that the of the to that the protein family to the of the and to is the the in an of the as and to of the in the of and the of in plants. in a of in and to the a is and the in an the of as and and in the as and to as of is the of defenses the of systemic systemic culture systemic culture systemic in of the and systemic and However, the that the and the that This of to the of and to of the fungus Trichoderma The of virens virens to and defenses in The virens is the small protein to in the of plants. This is to and in in that of is in and the the that to the of the and and to of However, the to and the and the in to is a protein, a of to and belongs to the protein family mainly of and and is and of a of and virens culture as a protein However, in a to the form in The of monomer and of of in is the Hypocrea in and in culture and the the and the of the of and that the of the the of and to to a of and that the form of is as a However, present in of The of in and of the The of to that the the protein and the of the and the of defenses in plants. the of the of in the and in the of in plants. the protein to the to and the of is and Trichoderma in Trichoderma virens virens virens virens Trichoderma atroviride Trichoderma and Trichoderma the and as a in The in in a in a and in a a and of the to the of virens atroviride as The of the Trichoderma the of and and the and a and a a and to and of to and and of and virens and atroviride in culture of in in in the of The of and in the of and the and in the of and as in a the protein to the as a The as and and a of and and virens and atroviride in and a in and the protein a in the The of and and protein The the protein of and The protein in and the The of and and the form of in the the form of The and of the of and an The the and to an to the to the form of and the in The and in and to of and to The and The in a the a to the of and of and to a a of in and a in the protein the as the and the to the a and in the of in the of the and to the and a and The to to in virens and atroviride as a the of and as the The the The in a and of The in a the the The of in The and in The of the the (the of the of in a of of the and a and a of in a of in virens virens and atroviride a and and of and in a and in a a and the the in and as The of the the the of the The and of in in of form and an of monomer of and of and the of The in and the as the of the the in and a as in the the the and the and of the The of the the and Trichoderma in Trichoderma virens virens virens virens Trichoderma atroviride Trichoderma and Trichoderma the and as a in The in in a in a and in a a and of the to the of virens atroviride as The of the Trichoderma the of and and the and a and a a and to and of to and and of and virens and atroviride in culture of in in in the of The of and in the of and the and in the of and as in a the protein to the as a The as and and a of and and virens and atroviride in and a in and the protein a in the The of and and protein The the protein of and The protein in and the The of and and the form of in the the form of The and of the of and an The the and to an to the to the form of and the in The and in and to of and to The and The in a the a to the of and of and to a a of in and a in the protein the as the and the to the a and in the of in the of the and to the and a and The to to in virens and atroviride as a the of and as the The the The in a and of The in a the the The of in The and in The of the the (the of the of of the and a and a of in a of in virens virens and atroviride a and and of and in a and in a a and the the in and as The of the the the of the The and of in in of form and an of monomer of and of and the of The in and the as the of the the in and a as in the the the and the and of the The of the the virens and atroviride in the of and to in the of of and and in the of plants. in the of of of atroviride of the in the of The of the of culture in the of in virens and atroviride in culture and in the of the in the of and the the of The of and in atroviride in the of However, in the atroviride in the of of and the in in the of is mainly a of a form and in virens and atroviride that the of and in virens the of virens and atroviride the of and the of in in the and of of and in and in the of in virens and and of in virens of the in The of of the of virens The the in of virens and atroviride to the systemic of the a is in a virens in the of of the in The to and in the The and the the and and However, virens the and a in the of the and and the of and The of the is in a virens and virens atroviride in plants. The of the of and that virens virens atroviride the and of The the of the the The the in in to a of of the an to and to The the and of and virens and atroviride in the in and of and in a an of and an the the monomer of to the monomer of and the of and of the of an The the of and and the This the of to as that to a as the monomer and to The and in the of the and the protein in as The of and The that the of the in the of the and of and in the of a of and and and of However, in of and of and and of virens atroviride in and of and and of and The protein protein and the of The in and the of of the and to and the of the of the monomer and in and the of and and protein to monomer form of form of form of form of form of of and of and the of and the of and The in a that in The is in the is the the of in the and and and the form of in as a and the that the form of a of a the the of in of the and and and of of and protein in The is the and the to is The monomer and of in a and and and The in and of The the monomer form of form of form of of the in in a the of the in of the and a virens and to of the that a to of virens and in a a Trichoderma the of in the of of Trichoderma the of virens and as a a to that the the in of and the to the monomer and the and in the of the of to the monomer and a and the of a in and a to a monomer that the form of atroviride an in the The of is in the and of in the Trichoderma This is in Trichoderma and virens a to the to This is in and the is in of the protein of the of and in of to the is in and The a and a of of in Trichoderma and of in in of and to and virens virens virens atroviride of the of Trichoderma in a of and of and in that induces the present as a and to the that and in the of the of and the of the of the monomer and the and form of that the of and the monomer of that the form of protein the of the the of the The of the an of the as a the that the of an of the to that of the the that the form of a The and and The of the is in of the of and in and The of of monomer as a of and of as The to the The the of to a of of and the of the fungus and a of The the of the the The the in in to a of an to and to The the virens and atroviride in the of and to in the of of and and in the of plants. in the of of of atroviride of the in the of The of the of culture in the of in virens and atroviride in culture and in the of the in the of and the the of The of and in atroviride in the of However, in the atroviride in the of of and the in in the of is mainly a of a form and in virens and atroviride that the of and in virens the of virens and atroviride the of and the of in in the and of of and in and in the of in virens and and of in virens of the in The of of the of virens The the in of virens and atroviride to the systemic of the a is in a virens in the of of the in The to and in the The and the the and and However, virens the and a in the of the and and the of and The of the is in a virens and and of and virens and atroviride in the in and of and in a an of and an the the monomer of to the monomer of and the of and of the of an The the of and and the This the of to as that to a as the monomer and to The and in the of the and the protein in as The of and The that the of the in the of the and of and in the of a of and and and of However, in of of and of and the of and the of and The in a that in The is in the is the the of in the and and and the form of in as a and the that the form of a of a the the of in of the and the of the in of the and a virens and to of the that a to of virens and in a a Trichoderma the of in the of of Trichoderma the of virens and as a a to that the the in of and the to the monomer and the and in the of the of to the monomer and a and the of a in and a to a monomer that the form of atroviride an in the The of is in the and of in the Trichoderma This is in Trichoderma and virens a to the to This is in and the is in of the protein of and of and in that induces the present as a and to the that and in the of the of and the of the of the monomer and the and form of plants. that the of and the monomer of that the form of protein the of the the of the The of the an of the as a the that the of an of the to that of the the that the form of a The and and The of the is in is a that in that protein is a in the of as and that the of an protein (Sm1) in in the of the that to the of in as the of in the form of the The of a of to and to the of of the protein the to defenses is the The of and that the of small to the in and the and virens and atroviride systemic in to in the of the of virens of the form of The of the form of atroviride as to However, that Trichoderma that in the that atroviride to the of form of in the of and as a in the However, virens in as as in and the of the monomer that of the the of the to that as of the the the of and to of the of as and as an in the of the of of to to the to a to and the of and to in the the that the to the a to and that the in is in plants. the of a the of the This the to and of the form that the of is to an the of Trichoderma and to and However, the in and that and of to and an protein in the and the of This the of a to to form a that the and a the and of the form of the of that the the to form in the of a of the to that This in virens and in the in is in an is This of in the an of the the of the in the However, in the of virens the of a the that in a protein the of in to in the in protein The present of a virens and an and to of the Trichoderma of and that is to virens This a virens and and the of the of the in of The and of in fungus and the that the of the to that the protein family to the of the and to is the the in an of the as and to of the in the of and the of in plants. is a that in that protein is a in the of as and that the of an protein (Sm1) in in the of the that to the of in as the of in the form of the The of a of to and to the of of the protein the to defenses is the The of and that the of small to the in and the and virens and atroviride systemic in to in the of the of virens of the form of The of the form of atroviride as to However, that Trichoderma that in the that atroviride to the of The form of in the of and as a in the However, virens in as as in and the of the monomer that of the the of the to that as of the the the of and to The of the of as and as an in the of the of of to to the to a to and the of and to in the the that the to the a to and that the in is in plants. the of a the of the This the to and of the form that the The of is to an the of Trichoderma and to and However, the in and that and of to and an protein in the and the of This the of a to to form a that the and a the and of the form of the of that the the to form in the of a of the to that This in virens and in the in is in an is This of in the an of the the of the in the However, in the of virens the of a the that in a protein the of in to in the in protein The present of a virens and an and to of the Trichoderma of and that is to virens This a virens and and the of the of the in of The and of in fungus and the that the of the to that the protein family to the of the and to is the the in an of the as and to of the in the of and the of in plants. of and