Physiology and Molecular Biology of Stress ... - KHAM PHA MOI
Physiology and Molecular Biology of Stress ... - KHAM PHA MOI
Physiology and Molecular Biology of Stress ... - KHAM PHA MOI
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144<br />
R.G. Trischuk, B.S. Schilling, M. Wisniewski <strong>and</strong> L.V. Gusta<br />
alcohols which increase in the autumn during acclimation <strong>and</strong> decrease in the spring<br />
during deacclimation (Levitt, 1980). Many wheat <strong>and</strong> canola cultivars show a parallel<br />
relationship between sugars <strong>and</strong> freezing tolerance. The main sugars that increase are<br />
sucrose, glucose, fructose, sorbitol, manitol, raffinose, <strong>and</strong> stachyose. It is postulated<br />
sugars replace water <strong>and</strong> decreases the degree <strong>of</strong> freeze-induced dehydration. Trehalose<br />
<strong>and</strong> umbelliferose have received considerable attention in the stress response as<br />
they promote glass transitions that protect cells from desiccation injury (Crowe et al.,<br />
1984, Wolkers et al., 1999). <strong>Stress</strong> induced proteins have a stabilizing effect on sugar<br />
glasses by increasing the average strength <strong>of</strong> hydrogen bonding in the dry state (Wolkers<br />
et al., 2001). Long chain fructans increase during cold acclimation <strong>of</strong> cereals <strong>and</strong> inhibit<br />
ice growth in xylem vessels (Olien, 1967). Thylakoid membranes are protected from<br />
freezing inactivation by exogenous proline, argenine, threonine <strong>and</strong> lysine. Proline <strong>and</strong><br />
glycine-betaine are both postulated to act as cryoprotectants. Yoshida <strong>and</strong> Uemura<br />
(1984) observed the development <strong>of</strong> freezing tolerance was accompanied by an increase<br />
in phospholipids, especially phosphatidyl choline <strong>and</strong> phosphatidyl ethanolamine.<br />
Free fatty acids were shown to accumulate as degradation products from ROS<br />
following a lethal-thaw event (McKersie <strong>and</strong> Bowley, 1996).<br />
Previously, it was impossible to measure all these changes simultaneously;<br />
however, ultra-high resolution mass spectrometry can identify <strong>and</strong> quantify over 100,000s<br />
<strong>of</strong> metabolites, simultaneously. The number <strong>of</strong> metabolites that can be identified <strong>and</strong><br />
quantified depends on the sample preparation x instrument sensitivity x physical concentration<br />
<strong>of</strong> the metabolite. Subtle differences in expressed genes <strong>and</strong> proteins can be<br />
verified through metabolomics.<br />
6. HORMONAL PROFILING<br />
In autumn, plants in the field are exposed to multiple stresses that play a role in determining<br />
their capacity to survive the winter. These stresses include low temperatures<br />
(both above <strong>and</strong> sub-zero), wind, drought, UV, photoinhibition, nutrients, salinity, high<br />
temperatures <strong>and</strong> mechanical injury. Plants use multiple signaling pathways <strong>and</strong> signals<br />
to mediate their acclimation responses. Two signaling pathways have been speculated<br />
to regulate cold acclimation (Thomashow, 1999); however, it is the specific combination<br />
<strong>of</strong> various components <strong>of</strong> the signaling network coupled with spatial <strong>and</strong> temporal<br />
factors that ultimately result in an increase in winter hardiness. Low temperature<br />
sensors may be due to changes in membrane fluidity (Murata <strong>and</strong> Los, 1997), conformational<br />
changes in proteins, altered ABA binding sites, release <strong>of</strong> sequestered ABA<br />
from plastids, decrease in cell water potential, etc. Secondary signals such as ABA <strong>and</strong><br />
second messengers can initiate a cascade <strong>of</strong> signaling events that may differ from the<br />
critical primary signal. For a comprehensive review on cell signaling the reader is<br />
referred to Xiong et al., (2002). Calcium has been demonstrated to act as a secondary<br />
messenger in low temperature signal transduction during cold acclimation (Monroy<br />
<strong>and</strong> Dhindsa, 1995). These studies are based on the observation <strong>of</strong> a transient increase