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Physiology and Molecular Biology of Stress ... - KHAM PHA MOI

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162<br />

A.R. Reddy <strong>and</strong> A.S. Raghavendra<br />

The ROS are also produced in different cellular components including chloroplasts,<br />

mitochondria, peroxisomes, glyoxysomes, cell wall, plasma membrane <strong>and</strong><br />

apoplasts (Figure 2). However, as depicted in Figure 2, the chloroplasts, mitochondria<br />

<strong>and</strong> the microbodies are the main sources <strong>of</strong> ROS in the plant cell.<br />

Figure 2. Generation <strong>of</strong> reactive oxygen species in different cellular compartments<br />

Although chloroplast was considered to be the main source <strong>of</strong> ROS production,<br />

recent studies suggest some intriguing possibilities about other cellular organelles<br />

as additional sources <strong>of</strong> ROS generation. Plant hypersensitive response <strong>and</strong> programmed<br />

cell death were partly attributed to the enhanced levels <strong>of</strong> ROS in mitochondria<br />

(Lam et al., 2001). In plant cells, mitochondrial ETC is a major site <strong>of</strong> ROS production<br />

(Moller, 2001; Tiwari et al., 2002). In addition to the complexes I-IV, the plant mitochondrial<br />

ETC contains proton pumping alternative oxidases as well as two non-proton<br />

pumping NAD(P)H dehydrogenases on each side <strong>of</strong> the inner membrane. Complex I is<br />

the main enzyme oxidizing NADH under normal conditions <strong>and</strong> is also a major site <strong>of</strong><br />

ROS generation (Figure 2). Several antioxidant enzymes are also reported in the matrix<br />

along with some antioxidants like glutathione to remove ROS produced under conditions<br />

<strong>of</strong> oxidative stress (Purvis, 1997; Braidot et al., 1999;). The entire ascorbate-glutathione<br />

cycle has been reported to occur in pea leaf mitochondria (Jimehez et al., 1998).

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