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

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276<br />

B. Rathinasabapathi <strong>and</strong> R. Kaur<br />

Rengel, 1999). Specific role <strong>of</strong> these enzymes or small molecules are better understood<br />

by manipulating expression <strong>of</strong> the genes encoding antioxidant enzymes or by engineering<br />

synthetic pathways to antioxidants (Allen et. al., 1997). Since many <strong>of</strong> both enzymatic<br />

<strong>and</strong> non-enzymatic antioxidant systems are understood in plants, it must be<br />

possible to improve oxidative stress tolerance in crops via metabolic engineering. Some<br />

examples for such metabolic engineering attempts are listed in Table 5.<br />

Transgenic tobacco, lucerne, potato <strong>and</strong> cotton overexpressing SOD in the<br />

chloroplasts showed higher tolerance to oxidative stress (Bowler et. al., 1991, Gupta et.<br />

al., 1993a, b, McKersie et. al., 1993, 1996, Perl et. al., 1993, Foyer et. al., 1994, Slooten et.<br />

al., 1995). Similar results were observed with overproduction <strong>of</strong> SOD in the mitochondria<br />

<strong>of</strong> lucerne (McKersie et. al., 1997) <strong>and</strong> in the cytosol <strong>of</strong> potato (Perl et. al., 1993).<br />

Transgenic tobacco engineered to overexpress MnSOD in chloroplast provided protection<br />

from Mn deficiency mediated oxidative stress (Yu et. al., 1999).<br />

The tripeptide glutathione, the major cellular antioxidant, detoxifies the excess<br />

hydrogen peroxide generated during oxidative stress. Glutathione metabolism has<br />

therefore been manipulated to improve plant’s oxidative stress tolerance (Noctor et. al.,<br />

1998). Transgenic plants overexpressing glutathione reductase had an increase in their<br />

reduced:oxidized glutathione ratio with enhanced tolerance to oxidative stress (Noctor<br />

et. al., 1998). Glutathione peroxidase is responsible for removal <strong>of</strong> unsaturated fatty acid<br />

hydroperoxides generated in cellular membranes during oxidative stress. Transgenic<br />

tobacco plants overexpressing Chlamydomonas glutathione peroxidase in the chloroplast<br />

or the cytosol, exhibited higher membrane integrity by increasing tolerance against<br />

oxidative stress (Yoshimura et. al., 2004).<br />

3.2.9. Heat <strong>and</strong> Cold <strong>Stress</strong><br />

Temperate <strong>and</strong> subtropical plants are extremely vulnerable to high temperatures especially<br />

during early tillering, flower initiation, anthesis <strong>and</strong> grain filling stages, leading to<br />

considerable decrease in crop productivity. All tropical crops are exposed to high temperatures<br />

<strong>and</strong> can benefit by increasing their high temperature stress tolerance further<br />

(also see Chapter 4). However, this trait is particularly valuable in some <strong>of</strong> the world’s<br />

most important food crops naturally adapted to cold weather. For example, improving<br />

the heat stress tolerance in winter cereals <strong>and</strong> potato can extend these crops to areas<br />

currently not suitable for them <strong>and</strong> increase their yield potential in some <strong>of</strong> the currently<br />

grown locations (Veilleux et. al., 1997; Maestri et. al., 2002). Despite our vast knowledge<br />

on how organisms respond to high temperature stress, little progress has been made in<br />

breeding crops for high temperature stress tolerance.<br />

3.2.10. Heat Shock Proteins<br />

In response to high temperature, all organisms including plants produce a set <strong>of</strong> proteins<br />

known as heat shock proteins (HSPs). Heat shock proteins (HSPs) have been

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