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Principles of terrestrial ecosystem ecology.pdf

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Leaf area (m 2 )<br />

500<br />

400<br />

300<br />

200<br />

100<br />

0 0<br />

Douglas fir<br />

Western white pine<br />

Ponderosa pine<br />

1000 2000 3000<br />

Sapwood area (cm 2 )<br />

Figure 4.13. Leaf area versus sapwood crosssectional<br />

area for three forest trees. Ponderosa pine,<br />

which typically occupies dry sites, has smaller vessels<br />

and therefore supports less leaf area per unit<br />

sapwood than does Douglas fir from moist sites.<br />

(Redrawn with permission from Canadian Journal <strong>of</strong><br />

Forest Research; Monserud and Marshall 1999.)<br />

conducting tissue (the sapwood). There is a<br />

strong linear relationship between the crosssectional<br />

area <strong>of</strong> sapwood and the leaf area supported<br />

by a tree (Fig. 4.13). However, the slope<br />

<strong>of</strong> this relationship varies strikingly among<br />

species and environments. Drought-resistant<br />

species generally have less leaf area per unit <strong>of</strong><br />

sapwood than do drought-sensitive species<br />

because <strong>of</strong> the small vessel diameter <strong>of</strong><br />

drought-resistant species. The ratio <strong>of</strong> leaf area<br />

to sapwood area, for example, is generally more<br />

Transpiration, (mL m -2 h -1 )<br />

30<br />

20<br />

10<br />

0<br />

0<br />

Transpiration<br />

Water Movements Within Ecosystems 87<br />

than twice as great in trees from mesic environments<br />

as in trees from dry environments,<br />

due to the smaller-diameter conducting elements<br />

in dry environments (Margolis et al.<br />

1995). Any factor that enhances the productivity<br />

<strong>of</strong> a tree increases its ratio <strong>of</strong> leaf area to<br />

sapwood area.This ratio increases, for example,<br />

with improvements in nutrient or moisture<br />

status and is greater in dominant than subdominant<br />

individuals <strong>of</strong> a stand.<br />

Water storage in stems buffers the plant from<br />

imbalances in water supply and demand. The<br />

water content <strong>of</strong> trunks <strong>of</strong> trees generally<br />

decreases during the day, causing water uptake<br />

by roots to lag behind transpirational water loss<br />

by about 2h (Fig. 4.14). The quantity <strong>of</strong> water<br />

stored in sapwood is substantial, equivalent to<br />

as much as 5 to 10 days <strong>of</strong> transpiration. This<br />

sapwood water, however, exchanges relatively<br />

slowly, so stores <strong>of</strong> water in sapwood seldom<br />

account for more than 10% <strong>of</strong> transpiration.<br />

In dry tropical forests, where trees lose their<br />

leaves during the dry season, this stored water<br />

is critical to support flowering during the dry<br />

season. Trees in these forests that have lowdensity<br />

wood and large stem water storage can<br />

flower during the dry season, whereas trees<br />

with high-density wood and low stem water<br />

storage can flower only during the wet season<br />

(Borchert 1994). Water storage in desert succulents<br />

may allow transpiration to continue for<br />

6 10 12<br />

Time (h)<br />

14 16 18 20 22 24 0<br />

2 4 8<br />

Figure 4.14. Diurnal time course <strong>of</strong> water uptake<br />

and water loss by Siberian larch. During morning,<br />

transpiration is supported by water loss from stems,<br />

creating a lower water potential in stems and roots,<br />

Water uptake<br />

10<br />

5<br />

Water uptake (L h -1 )<br />

which generates the water potential gradient to<br />

absorb water from the soil.The water stored in stems<br />

is replenished at night. (Redrawn with permission<br />

from BioScience; Schulze et al. 1987.)

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