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

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112<br />

T.D. Sharkey <strong>and</strong> S.M. Schrader<br />

<strong>of</strong> an increased pH gradient (Horton et al., 1996). Further, ATP levels, which are dependent<br />

on an energized thylakoid membrane, were higher in isolated chloroplasts (Weis,<br />

1981a) <strong>and</strong> did not change in whole leaves (Schrader et al., 2004) under heat stress.<br />

3.3.3. The Role <strong>of</strong> Membrane-Bound Reactions<br />

Early studies <strong>of</strong> high temperature stress on photosynthesis noted a decline in oxygen<br />

evolution in isolated chloroplasts (Holt <strong>and</strong> French, 1946). Katoh <strong>and</strong> San Pietro (1967)<br />

ascribed this heat effect on O 2<br />

evolution to the water splitting complex <strong>of</strong> PSII, showing<br />

that NADP photoreduction could be restored using ascorbate as an artificial electron<br />

donor that substituted for the water splitting complex. Later, the heat inactivation <strong>of</strong> O 2<br />

evolution was associated with the release <strong>of</strong> Mn from PSII (Cheniae <strong>and</strong> Martin, 1970;<br />

Kimimura <strong>and</strong> Katoh, 1972; Wydrzynski <strong>and</strong> Sauer, 1980) <strong>and</strong> the release <strong>of</strong> the 18, 24,<br />

<strong>and</strong> 33 kDa proteins from the thylakoid membranes which are associated with the water<br />

splitting complex on the luminal surface <strong>of</strong> PSII (Figure 2) (Yamamoto <strong>and</strong> Nishimura,<br />

1983; Nash et al., 1985).<br />

Figure 2. Photosynthetic electron transport <strong>and</strong> processes that can be affected by high<br />

temperature stress (Light arrows indicate electron transport paths, heavy arrows indicate<br />

changes that can occur in response to heat)

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