Agriculture Reference
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In addition, another study has shown quercetin, which is a phenolic abundant in apples,
to be an aldose reductase (alditol: NADP
oxidoreductase) inhibitor (Costantino et al.,
1999). Aldose reductase is the first enzyme of the polyol pathway. Glucose metabolism
through the polyol pathway has been linked to long-term diabetic complications like cataract,
nephropathy, neuropathy, and retinopathy (Costantino et al., 1999).
+
16.4 Role of free radical-scavenging and enzyme antioxidant
activity in postharvest preservation of fruits
Reactive oxygen species (ROS) like superoxide radicals (O
.
2
), hydroxyl radicals (
.
OH),
and hydrogen peroxide (H
2
O
2
) are the products of oxidative dysfunctional biochemical
reactions within cells. These ROS when left unchecked cause oxidative damage, resulting
in lipid peroxidation, protein denaturation, and mutagenesis. An increase in ROS is linked
to different types of stress such as drought, heat stress, metal toxicity, radiation exposure,
pathogens, and salinity (Bolwell and Wojitaszek, 1997; Jimenez et al., 1998; Karpinski,
et al., 1999; Dat et al., 2000; Hernandez et al., 2001; Quartacci et al., 2001; Del Rio et al.,
2002; Fig. 16.4). Further ROS is also involved in natural and induced senescence and cell
death in plants (Droillard et al., 1987; Thompson et al., 1991; Philosoph-Hadas et al., 1994;
Bartoli et al., 1996). For example, studies have indicated increase in hydroperoxides during
pepper, banana, pear, and tomato ripening during which senescence is induced (Frenkel,
1978; Thompson et al., 1987; Rogiers et al., 1998).
Plants counter harmful effects of ROS with antioxidants metabolites and enzymes. An-
tioxidants metabolites include water-soluble compounds like ascorbate, glutathione, and
flavonoids and lipid-soluble compounds like carotenoids and tocopherols. Enzymes linked
to antioxidant response include superoxide dismutase (SOD), catalase (CAT), monodehy-
droascorbate reductase (MDHAR), dehydroascorbate reductase (DHAR), ascorbate per-
oxidase (ASPX), and glutathione reductase (GR). SOD converts O
.
2
to H
2
O
2
, which is
then reduced to H
2
O by CAT or ASPX-depending cellular localization (Foyer et al., 1994;
Hodges et al., 1996; Hodges and Forney, 2000; Fig. 16.5). Both ascorbate and glutathione
can also interact directly with and scavenge ROS (Foyer et al., 1994). Oxidized ascorbate
is reduced by DHAR, MDHAR, and glutathione, and oxidized glutathione is reduced by
GR. Reduction of both oxidized glutathione and ascorbate by their respective enzymes
is NADPH dependant (Shetty and Wahlqvist, 2004). Studies have shown many phenolic
compounds especially flavonoids, for example, quercetin, rutin, and catechin have free
radical-scavenging antioxidant activity (Kang et al., 2004; Zhang et al., 2006). In addition,
studies in fava beans and peas have shown that even exogenous phenolics can stimulate
antioxidant enzyme activity (Duval and Shetty, 2000; Vattem et al., 2005).
Therefore, since ROS is involved in plant development, including fruit ripening and
senescence, antioxidant response coupling phenolic synthesis and antioxidant enzyme re-
sponse may be recruited to counter ROS and senescence (Pastori and Del-Rio, 1997;
Jimenez et al., 1998, 2002, 2003; Hodges and Forney, 2000). Previous studies with
muskmelon fruits and sunflower seeds indicated that delayed senescence in specific tis-
sue types correlated to high antioxidant enzyme response (Bailly et al., 1996; Lacan and
Baccou, 1998). In order to couple cellular antioxidants like ascorbate, glutathione and phe-
nolic phytochemicals with antioxidant enzymes for effective antioxidant response cellular
reducing equivalents such as FADH
2
and NADPH are required. Studies have found that
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