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Department of Tumor Immunology, University Medical Center Nijmegen, Nijmegen, The Netherlands (F.A.D.T.G.W., G.J.A., C.G.F.); Department of Medical Immunology, Charité, Humboldt University, Berlin, Germany (H.-D.V.); Department of Pharmacology, University College, London, England (D.W.); Department of Pharmacology, New York Medical College, Valhalla, New York (N.G.A.); Beth Israel Deaconess Medical Center, Department of Surgery, Harvard Medical School, Boston, Massachusetts (M.P.S.); and Inflammation Laboratory, Institute Gulbenkian for Science, Lisbon, Portugal (M.P.S.)
The heme-heme oxygenase system has recently been recognized to possess important regulatory properties. It is tightly involved in both physiological as well as pathophysiological processes, such as cytoprotection, apoptosis, and inflammation. Heme functions as a double-edged sword. In moderate quantities and bound to protein, it forms an essential element for various biological processes, but when unleashed in large amounts, it can become toxic by mediating oxidative stress and inflammation. The effect of this free heme on the vascular system is determined by extracellular factors, such as hemoglobin/heme-binding proteins, haptoglobin, albumin, and hemopexin, and intracellular factors, including heme oxygenases and ferritin. Heme oxygenase (HO) enzyme activity results in the degradation of heme and the production of iron, carbon monoxide, and biliverdin. All these heme-degradation products are potentially toxic, but may also provide strong cytoprotection, depending on the generated amounts and the microenvironment. Pre-induction of HO activity has been demonstrated to ameliorate inflammation and mediate potent resistance to oxidative injury. A better understanding of the complex heme-heme
Abstract I. Introduction II. Heme A. Heme Biosynthesis B. Heme As Active Site in Heme Proteins C. Heme-Mediated Gene Modulation, Cell Differentiation, Proliferation, and Immune Stimulation D. Toxic Effects of Free Heme and Its Control III. Heme Oxygenase: The Heme-Degrading Enzyme A. History of Heme Oxygenase B. Heme Oxygenase Isoforms and Gene Regulation C. Traditional View on Heme Oxygenase and Its Toxic Breakdown Products D. Recent Insights in Heme Oxygenase and a Novel Role for Heme-Derived Metabolites IV. Inflammation and the Role of Adhesion Molecules A. The Role of Heme in Inflammation B. Consequences of Heme-Induced Oxidative Stress and Inflammation C. The Role of Heme Oxygenase in Inflammation D. Mechanism of Heme Oxygenase-Mediated Down-Modulation of Inflammation E. Effects of Heme Oxygenase Inhibition versus Heme Oxygenase-1 Overexpression V. The Heme-Heme Oxygenase System A. The Heme-Heme Oxygenase-1 System and Apoptosis B. The Heme-Heme Oxygenase System and Transplantation Biology C. The Heme-Heme Oxygenase System As Target for Inflammatory Control D. Some Considerations Concerning Modulation of the Heme-Heme Oxygenase System VI. Conclusion
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