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Background  The adage from Shakespeare, "troubles, not as single spies, but in battalions come," holds true for  Nicotiana attenuata , which is commonly attacked by both pathogens ( Pseudomonas spp .) and herbivores ( Manduca sexta ) in its native habitats. Defense responses targeted against the pathogens can directly or indirectly influence the responses against the herbivores. Na defensin  is an effective induced defense gene against the bacterial pathogen  Pseudomonas syringae  pv  tomato  (PST DC3000), which is also elicited by attack from  M. sexta  larvae, but whether this defense protein influences  M. sexta's  growth and whether  M. sexta -induced Na defensin  directly or indirectly influences PST DC3000 resistance are unknown.    Results   M. sexta  larvae consumed less on WT and on Na defensin -silenced  N. attenuata  plants that had previously been infected with PST DC3000 than on uninfected plants. WT plants infected with PST DC3000 showed enhanced resistance to PST DC3000 and decreased leaf consumption by  M. sexta  larvae, but larval mass gain was unaffected. PST DC3000-infected Na defensin -silenced plants were less resistant to subsequent PST DC3000 challenge, and on these plants,  M. sexta  larvae consumed less and gained less mass. WT and Na defensin -silenced plants previously damaged by  M. sexta  larvae were better able to resist subsequent PST DC3000 challenges than were undamaged plants.    Conclusion  These results demonstrate that Na-defensin directly mediates defense against PST DC3000 and indirectly against  M. sexta  in  N. attenuata . In plants that were previously infected with PST DC3000, the altered leaf chemistry in PST DC3000-resistant WT plants and PST DC3000-susceptible Na defensin -silenced plants differentially reduced  M. sexta's  leaf consumption and mass gain. In plants that were previously damaged by  M. sexta , the combined effect of the altered host plant chemistry and a broad spectrum of anti-herbivore induced metabolomic responses was more effective than Na defensin  alone in resisting PST DC3000.

Host-plant-mediated effects of Nadefensin on herbivore and pathogen resistance in Nicotiana attenuata