intense exercise alters the balance between oxidants/antioxidants and mitochondrial respiratory enzymes in the skeletal muscle.

In rodents, moderate exercise training ( swimming) has been shown to reduce the basal rate of mitochondrial hydrogen peroxide (H2O2) production (Venditti et al. 1999) and to enhance antioxidant enzyme activity, adaptations that would expected to reduce tissue oxidative damage (Leeuwenburgh et al. 1997). Venditti P, Masullo P, Di Meo S (1999) EVect of training on H2O2 Leeuwenburgh C, Wagner P, Holloszy JO, Sohal RS, Heinecke JW (1997) Caloric restriction attenuates dityrosine cross-linking of cardiac and skeletal muscle proteins in aging mice. Arch Biochem Biophys 346:7480. doi:10.1006/abbi.1997.0297 release by mitochondria from rat skeletal muscle. Arch Biochem

Free radicals can be defined as molecules or molecular fragments containing one or more unpaired electronsin atomic or molecular orbitals (Halliwell&Gutteridge, 1999). This unpaired electron(s) usually gives a considerable degree of reactivity to the free radical. Radicals derived from oxygen represent the most important class of radical species generated in living systems (Miller, Buettner, &Aust, 1990).

Miller, E. R., Pastor-Barriuso, R., Dalal, D., Riemersma, R. A., Appel, L. J., &Guallar, E. (2005). Meta-analysis: High-dosage Vitamin E supplementation may increase all-cause mortality. Ann. Intern. Med., 142, 3746.Valko, M.,Morris, H., & Cronin, M. T. D. (2005). Metals, toxicity and oxidative stress. Curr. Med. Chem., 12, 11611208.

Antioxidant enzymes Changes in antioxidant enzyme activity in erythrocytes have been used to document oxidative stress. The enzymes that have been most commonly examined after exercise stress are superoxide dismutase (SOD), catalase, glutathione peroxidase,and glutathione reductase. Maria L. Urso *, Priscilla M. Clarkson

Exercise Science Department, University of Massachusetts, 110 Totman Building, Amherst, MA 01003, USA (2003)

A Superoxide dismutase (SOD), which serves as a scavenger of 0;) is one of the most important enzymes in the antioxidant defense system. Mammalian tissues contain two main forms of SOD: Mn-SOD is present mostly in the mitochondrial matrix, whereas Cu,ZnSOD is predominantly localized in the cytosol. Since Tolmasoff et al. [12] pointed out that the ratio of SOD specific activity to specific metabolic rate of the tissue or of the whole adult organism increases with increasing maximum life-span potential for all the species, J.M. Tolmasoff, T. Ono and R.G. Cutler, Superoxide dismutase: correlation with life-span and specific metabolic rate in primate species. Proc. Natl. Acad. Sci. USA, 77 (1980) 2777-2781.

Lipid peroxidation Measures of lipid peroxidation include expired pentane, malondialdehydes (MDA), lipid hydroperoxides, isoprostanes, and conjugated dienes. Most studies have used MDA as a measure of oxidative stress imposed by exercise. When free radicals are generated they can attack polyunsaturated fatty acids in the cell membrane leading to a chain of chemical reactions called lipid peroxidation. As the fatty acid is broken down, hydrocarbon gases (ethane or pentane) and aldehydes are formed. Maria L. Urso *, Priscilla M. Clarkson Exercise Science Department, University of Massachusetts, 110 Totman Building, Amherst, MA 01003, USA

Malondialdehyde Aldehydes, especially MDA, have been frequently used as markers of oxidative stress in response to exercise.(Halliwell and Chirico, 1993; HanEt al., 2000). Halliwell, B., Chirico, S., 1993. Lipid peroxidation: its mechanism, measurement, and significance. Am. J. Clin. Nutr. 57 (5), 715S_/724. Han, D., Loukianoff, S., McLaughlin, L., 2000. In: Hanninen,

O., Packer, L., Sen, C.K. (Eds.), Handbook of Oxidants and Antioxidants in Exercise. Elsevier, Amsterdam, pp. 433_/ 484\