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

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104<br />

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

cotton, which normally grows in warm environments, stomatal conductance can be so<br />

high that leaf temperature is substantially below air temperature. In fact, it has been<br />

specifically shown for cotton that the greater the transpirational cooling, the greater the<br />

yield (Radin et al., 1994; Lu et al., 1997) proving that heat-stress-induced reductions <strong>of</strong><br />

photosynthesis limit overall yield. Other studies have also linked reduced crop yields<br />

to moderate heat stress <strong>of</strong> leaves that reduces photosynthesis (Al-Khatib <strong>and</strong> Paulsen,<br />

1990; Guilioni et al., 2003).<br />

Some important effects <strong>of</strong> energy balance can be seen in the oak leaf temperatures<br />

shown in Figure 1. During the day, leaves at the top <strong>of</strong> the tree (black dots) can be<br />

over 10°C above air temperature because <strong>of</strong> the radiant heat load, while leaves at the<br />

bottom <strong>of</strong> the tree (light gray dots) are slightly below air temperature because <strong>of</strong> latent<br />

heat loss. At night, leaves at the top <strong>of</strong> the tree lose radiant heat to the sky <strong>and</strong> so are<br />

cooler than leaves at the bottom <strong>of</strong> the tree because leaves at the bottom <strong>of</strong> the tree<br />

exchange radiant heat with other leaves <strong>and</strong> with the ground, both <strong>of</strong> which are warmer<br />

than the sky.<br />

12<br />

Leaf Temp - Air Temp, o C<br />

10<br />

8<br />

6<br />

4<br />

2<br />

0<br />

- 2<br />

- 4<br />

10:00 20:00 6:00 16:00 2:00 12:00<br />

Time, Eastern Daylight Time starting July 10<br />

Figure 1. Difference between leaf <strong>and</strong> air temperature for two days for three oak (Quercus<br />

rubra) leaves at the top (black dots) middle (dark gray) or bottom (light gray) <strong>of</strong> the canopy<br />

Leaves exposed to full sunlight will carry out most photosynthesis but also be<br />

subject to the rapid, high temperature spikes shown in Figure 1. We have started calling<br />

this heat flecks, analogous to (<strong>and</strong> sometimes simultaneous with) sunflecks (Pearcy et<br />

al., 1996). Because leaves are <strong>of</strong>ten the hottest parts <strong>of</strong> plants, high temperature stress<br />

is especially relevant to photosynthesis. Moreover, plant growth <strong>of</strong>ten increases with<br />

temperature to well above 30°C while photosynthesis <strong>of</strong> individual leaves typically has<br />

a temperature optimum <strong>of</strong> 30°C or less. At low temperature growth may be more directly

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