Antioxidants in Health and Disease J Clin Pathol: First Published As 10.1136/Jcp.54.3.176 on 1 March 2001

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Antioxidants in Health and Disease J Clin Pathol: First Published As 10.1136/Jcp.54.3.176 on 1 March 2001 176 J Clin Pathol 2001;54:176–186 Antioxidants in health and disease J Clin Pathol: first published as 10.1136/jcp.54.3.176 on 1 March 2001. Downloaded from I S Young, J V Woodside Abstract induced tissue damage, and the function of Free radical production occurs continu- antioxidant defence systems in health and dis- ously in all cells as part of normal cellular ease. function. However, excess free radical production originating from endogenous Free radicals and their chemical or exogenous sources might play a role in reactions many diseases. Antioxidants prevent free A free radical can be defined as any molecular radical induced tissue damage by prevent- species capable of independent existence that ing the formation of radicals, scavenging contains an unpaired electron in an atomic them, or by promoting their decomposi- orbital.2 The presence of an unpaired electron tion. This article reviews the basic chem- results in certain common properties that are istry of free radical formation in the body, shared by most radicals. Radicals are weakly the consequences of free radical induced attracted to a magnetic field and are said to be tissue damage, and the function of anti- paramagnetic. Many radicals are highly reac- oxidant defence systems, with particular tive and can either donate an electron to or reference to the development of athero- extract an electron from other molecules, sclerosis. therefore behaving as oxidants or reductants. Department of Clinical (J Clin Pathol 2001;54:176–186) As a result of this high reactivity, most radicals Biochemistry, Institute have a very short half life (10−6 seconds or less) of Clinical Science, Keywords: free radicals; antioxidants; oxidative stress; coronary heart disease; atherosclerosis in biological systems, although some species Grosvenor Road, may survive for much longer.2 The most Belfast, Northern Ireland, BT12 6BJ, UK important free radicals in many disease states I S Young An antioxidant can be defined as: “any are oxygen derivatives, particularly superoxide substance that, when present in low concentra- and the hydroxyl radical. Radical formation in Department of tions compared to that of an oxidisable the body occurs by several mechanisms, Surgery, Royal Free involving both endogenous and environmental substrate, significantly delays or inhibits the and University College factors (fig 1). London Medical oxidation of that substrate”.1 The physiological Superoxide (O −.) is produced by the addi- School, 67–73 Riding role of antioxidants, as this definition suggests, 2 tion of a single electron to oxygen, and several House Street, London, is to prevent damage to cellular components W1P 7LD, UK mechanisms exist by which superoxide can be arising as a consequence of chemical reactions 3 J V Woodside produced in vivo. Several molecules, including http://jcp.bmj.com/ involving free radicals. In recent years, a adrenaline, flavine nucleotides, thiol com- Correspondence to: substantial body of evidence has developed Professor Young, pounds, and glucose, can oxidise in the Department of Clinical supporting a key role for free radicals in many presence of oxygen to produce superoxide, and Biochemistry, Institute of fundamental cellular reactions and suggesting these reactions are greatly accelerated by the Clinical Science, Royal that oxidative stress might be important in the Group of Hospitals, presence of transition metals such as iron or Grosvenor Road, Belfast pathophysiology of common diseases including copper. The electron transport chain in the BT12 6BJ, UK atherosclerosis, chronic renal failure, and inner mitochondrial membrane performs the [email protected] diabetes mellitus. The aim of this review is to reduction of oxygen to water. During this on October 1, 2021 by guest. Protected copyright. Accepted for publication consider mechanisms of free radical formation process free radical intermediates are gener- 5 June 2000 in the body, the consequences of free radical ated, which are generally tightly bound to the components of the transport chain. However, Endogenous sources Environmental sources Free radical production there is a constant leak of a few electrons into •mitochondrial leak •cigarette smoke the mitochondrial matrix and this results in the •respiratory burst •pollutants 4 •enzyme reactions •UV light formation of superoxide. The activity of O –., H O •autooxidation reactions 2 2 2 •ionising radiation several other enzymes, such as cytochrome •xenobiotics p450 oxidase in the liver and enzymes involved Transition in the synthesis of adrenal hormones, also metals results in the leakage of a few electrons into the Fe2+, Cu+ surrounding cytoplasm and hence superoxide formation. There might also be continuous OH. production of superoxide by vascular endothe- lium to neutralise nitric oxide,56production of superoxide by other cells to regulate cell growth and diVerentiation,7 and the produc- Lipid peroxidationModified DNA bases Protein damage tion of superoxide by phagocytic cells during the respiratory burst.8 Any biological system generating superoxide will also produce hydrogen peroxide as a result Tissue damage of a spontaneous dismutation reaction. In Figure 1 Major sources of free radicals in the body and the consequences of free radical addition, several enzymatic reactions, includ- damage. ing those catalysed by glycolate oxidase and www.jclinpath.com Antioxidants in health and disease 177 D-amino acid oxidase, might produce hydro- The net result of the reaction sequence illus- gen peroxide directly.9 Hydrogen peroxide is trated above is known as the Haber-Weiss reac- not a free radical itself, but is usually included tion. Although most iron and copper in the J Clin Pathol: first published as 10.1136/jcp.54.3.176 on 1 March 2001. Downloaded from under the general heading of reactive oxygen body are sequestered in forms that are not species (ROS). It is a weak oxidising agent that available to catalyse this reaction sequence, it is might directly damage proteins and enzymes still of importance as a mechanism for the for- containing reactive thiol groups. However, its mation of the hydroxyl radical in vivo. The most vital property is the ability to cross cell actual reactions, however, may be somewhat membranes freely, which superoxide generally more complex than those described above and cannot do.10 Therefore, hydrogen peroxide it is possible that other reactive intermediates such as the ferryl and perferryl radicals might formed in one location might diVuse a consid- 12 erable distance before decomposing to yield also be formed. the highly reactive hydroxyl radical, which is Approximately 4.5 g of iron can be found in the average adult man, most of which is likely to mediate most of the toxic eVects contained in the haemoglobin molecule and ascribed to hydrogen peroxide. Therefore, other haem containing proteins. Dietary iron is hydrogen peroxide acts as a conduit to transmit absorbed preferentially from the proximal part free radical induced damage across cell com- of the small intestine in the divalent form and is partments and between cells. In the presence of transferred to the circulation in which it is car- hydrogen peroxide, myeloperoxidase will gen- ried by transferrin.14 Under most circum- erate hypochlorous acid and singlet oxygen, a stances iron remains tightly bound to one of reaction that plays an important role in the 11 several proteins, including transferrin, lactofer- killing of bacteria by phagocytes. rin, haem proteins, ferritin, or haemosiderin. In . The hydroxyl radical (OH ), or a closely addition, however, it seems likely that a small related species, is probably the final mediator of iron pool will be maintained as complexes with 12 most free radical induced tissue damage. All a variety of small molecules, such as nucleo- of the reactive oxygen species described above tides and citrate within the cytoplasm and exert most of their pathological eVects by subcellular organelles.14 This pool is probably giving rise to hydroxyl radical formation. The capable of catalysing an iron driven Fenton reason for this is that the hydroxyl radical reaction in vivo. Certainly, these complexes can reacts, with extremely high rate constants, with promote hydroxyl radical formation in vitro.15 almost every type of molecule found in living Redox reactive iron can be measured using the cells including sugars, amino acids, lipids, and bleomycin iron assay,16 although it remains nucleotides. Although hydroxyl radical forma- unclear to what extent iron detected by this tion can occur in several ways, by far the most assay correlates with any discrete anatomical or important mechanism in vivo is likely to be the physiological pool. In normal circumstances, transition metal catalysed decomposition of no bleomycin reactive iron is detectable in superoxide and hydrogen peroxide.13 plasma from healthy subjects, implying that All elements in the first row of the d-block of transferrin or ferritin bound iron is not http://jcp.bmj.com/ the periodic table are classified as transition available to drive hydroxyl radical production.17 metals. In general, they contain one or more However, transferrin will release its iron at an unpaired electrons and are therefore them- acidic pH, particularly in the presence of small selves radicals when in the elemental state. molecular weight chelating agents such as 15 However, their key property from the point of ADP, ATP, and citrate. Such conditions are view of free radical biology is their variable found in areas of active inflammation
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