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352                Natural Antioxidants: Applications in Foods of Animal Origin
  VetBooks.ir  9   compatibility with the food and ease of application



               9   survive after processing and be stable in the finished product
               9   available at low cost.

               The compound and its oxidation products must also be nontoxic, even at
            doses much larger than those that normally would be ingested in food.
               Antioxidants can be classified according to the mechanism of action into
            two groups:



            9.2.1.4.1  Primary Antioxidants

            Primary antioxidants interfere with autoxidation by interrupting the chain
            propagation  mechanism.  Primary antioxidants  are free radical  acceptors
            (Wasowicz et al., 2004). The best-known and most effective primary anti-
            oxidant substances are polyphenols. They react with the chain-propagating
            radical species, which results in the formation of radical species incapable of
            extracting hydrogen atoms from unsaturated lipids (Coma & Kerry, 2012).
               Tocopherols inhibit  lipid oxidation by scavenging of aqueous and
            lipophilic free radicals as well as physical effects on membrane structure
            (Buettner, 1993; Atkinson et al., 2008). Dietary antioxidant treatments (i.e.,
            the  inclusion  of  antioxidants  in  animal  feed)  have  been  shown to  stabi-
            lize lipids in membranes and reduce the extent of lipid oxidation in meat
            during storage, but antioxidant effects in meat can differ between muscle
            types (Morrissey et al., 1997; Ahn et al., 2006). Vitamin E in livestock diets
            has been shown to reduce lipid oxidation in meats (Morrissey et al., 1998;
            Álvarez et al., 2009). Batifoulier et al. (2002) reported that supplementation
            of turkeys with α-tocopheryl acetate increased vitamin E content of micro-
            somal membranes and had also a protective effect on lipid oxidation.
               Some endogenous enzymes also have an antioxidant effect such as super-
            oxide dismutase, catalase or glutathione peroxidase. These enzymes inhibit
            lipid oxidation through the following mechanisms:

               -   Superoxide  dismutase  is present  in cells  and  extracellular fluids
                   to remove  •O  resulting in formation of oxygen and hydrogen
                             −
                                2
                   peroxide.
               -   Catalase, a heme-containing enzyme reacts with H O  to form water
                                                                 2
                                                               2
                   and oxygen (Goth, 1987; Richards, 2006).
               -   Glutathione peroxidase reduces hydrogen peroxide and lipid
                   hydroperoxides to alcohols (Gong et al., 2010).
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