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B 1
* Division of Pulmonary Sciences and Critical Care Medicine, University of Colorado Health Sciences Center, Denver, CO 80262; and
Service d’ Anesthésie-Reanimation et Unité Propre de Recherche de l’ Enseignement Superieur-Equipe d’ Accueil, Hôpital de Bicêtre, Le Kremlin Bicetre, France
Although oxidative stress has been thought to play a general role in the activation of NF-
B, the involvement of reactive oxygen species (ROS) in facilitating nuclear translocation of NF-B in neutrophils has not been described. In addition, the mechanisms by which ROS modulate the transcriptional activity of NF-B in response to Toll-like receptor 4 (TLR4)-dependent signaling are not well characterized. To examine these issues, oxidant-dependent signaling events downstream of TLR4 were investigated in neutrophils stimulated with LPS. Pretreatment of neutrophils with the antioxidants N-acetylcysteine or 
-tocopherol prevented LPS-induced nuclear translocation of NF-B. Antioxidant treatment of LPS-stimulated neutrophils also inhibited the production of proinflammatory cytokines (TNF-, macrophage inflammatory protein-2, and IL-1
), as well as activation of the kinases IB kinase , IB kinase , p38, Akt, and extracellular receptor-activated kinases 1 and 2. The decrease in cytoplasmic levels of IB produced by exposure of neutrophils to LPS was prevented by N-acetylcysteine or -tocopherol. Activation of IL-1R-associated kinase-1 (IRAK-1) and IRAK-4 in response to LPS stimulation was inhibited by antioxidants. These results demonstrate that proximal events in TLR4 signaling, at or antecedent to IRAK-1 and IRAK-4 activation, are oxidant dependent and indicate that ROS can modulate NF-B-dependent transcription through their involvement in early TLR4-mediated cellular responses.
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