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Background  Mitogen-activated protein kinase (MAPK) cascades are highly conserved signaling modules that mediate the transduction of extracellular stimuli via receptors/sensors into intracellular responses and play key roles in plant immunity against pathogen attack. However, the function of tomato MAPK kinases, SlMKKs, in resistance against  Botrytis cinerea  remains unclear yet.    Results  A total of five  SlMKK  genes with one new member,  SlMKK5 , were identified in tomato. qRT-PCR analyses revealed that expression of  SlMKK2  and  SlMKK4  was strongly induced by  B. cinerea  and by jasmonic acid and ethylene precursor 1-amino cyclopropane-1-carboxylic acid. Virus-induced gene silencing (VIGS)-based knockdown of individual  SlMKKs  and disease assays identified that  SlMKK2  and  SlMKK4  but not other three  SlMKKs  ( SlMKK1 ,  SlMKK3  and  SlMKK5 ) are involved in resistance against  B. cinerea . Silencing of  SlMKK2  or  SlMKK4  resulted in reduced resistance to  B. cinerea , increased accumulation of reactive oxygen species and attenuated expression of defense genes after infection of  B. cinerea  in tomato plants. Furthermore, transient expression of constitutively active phosphomimicking forms  SlMKK2   DD   and  SlMKK4   DD   in leaves of  Nicotiana benthamiana  plants led to enhanced resistance to  B. cinerea  and elevated expression of defense genes.    Conclusions  VIGS-based knockdown of  SlMKK2  and  SlMKK4  expression in tomato and gain-of-function transient expression of constitutively active phosphomimicking forms  SlMKK2   DD   and  SlMKK2   DD   in  N. benthamiana  demonstrate that both SlMKK2 and SlMKK4 function as positive regulators of defense response against  B. cinerea .

Tomato SlMKK2 and SlMKK4 contribute to disease resistance against Botrytis cinerea