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Principles of terrestrial ecosystem ecology.pdf

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Depth (m)<br />

0<br />

0.2<br />

0.4<br />

0.6<br />

0.8<br />

1.0<br />

1.2<br />

1.4<br />

Chaparral<br />

1.6<br />

ψplant Quercus -0.2<br />

1.8<br />

Adenostoma -1.4<br />

Heteromeles -2.1<br />

2.0 Rhamnus -2.8<br />

Grassland<br />

ψplant Sitanion < -4.0<br />

-5.0 -4.0 -3.0 -2..0 -1.0 0<br />

Soil water potential , ψ soil (MPa)<br />

Figure 4.8. Soil water pr<strong>of</strong>iles in adjacent shrub and<br />

grassland communities at the end <strong>of</strong> the summer<br />

drought period. Predawn water potentials are a good<br />

index <strong>of</strong> the soil moisture and the degree <strong>of</strong> drought<br />

stress experienced by the plant. (Redrawn with permission<br />

from Oecologia; Davis and Mooney 1986.)<br />

Water Movement Through Plants<br />

The vapor-pressure gradient from the leaf<br />

surface to the atmosphere is the driving force<br />

for water movement through plants. Water<br />

transport from the soil through the plant to<br />

the atmosphere takes place in a soil-plantatmosphere<br />

continuum that is interconnected<br />

by a continuous film <strong>of</strong> liquid water. Water<br />

moves from the soil through the plant to the<br />

atmosphere along a gradient in water potential.<br />

The low partial pressure <strong>of</strong> water vapor in air<br />

relative to that inside the leaves is the major<br />

Water Movements Within Ecosystems 83<br />

driving force for water loss from leaves, which<br />

in turn drives water transport along a pressure<br />

gradient from the roots to the leaves, which in<br />

turn drives water movement from the soil into<br />

the plant. The rate <strong>of</strong> water movement through<br />

the plant (J p) (Eq. 4.7) is determined by the<br />

water-potential gradient (the driving force;<br />

DYt) and the resistance to water movement, just<br />

as described for water movement through soils<br />

(Eq. 4.6). As in soils, the resistance to water<br />

movement through the plant depends on<br />

hydraulic conductivity (Lp) and path length (l).<br />

(4.7)<br />

The movement <strong>of</strong> water into and through the<br />

plant is driven entirely by the physical process<br />

<strong>of</strong> evaporation from the leaf surface and involves<br />

no expenditure <strong>of</strong> metabolic energy by<br />

the plant. This contrasts with the acquisition <strong>of</strong><br />

carbon and nutrients for which the plant must<br />

expend considerable metabolic energy (see<br />

Chapters 5 and 8).<br />

Roots<br />

DYt<br />

Jp = Lp l<br />

Water moves through roots along a waterpotential<br />

gradient from moist soils to the<br />

atmosphere during the day and sometimes to<br />

dry surface soils at night. In moist soils, the cell<br />

membranes, which are composed <strong>of</strong> hydrophobic<br />

lipids, provide the greatest resistance to<br />

water movement through roots (see Fig. 8.3).<br />

This membrane resistance to water flow is<br />

greatest under conditions <strong>of</strong> low root temperature<br />

or flooding, so plants that are not adapted<br />

to these conditions experience substantial<br />

water stress in cold or saturated soils. In dry<br />

soils the contact between the root and the<br />

soil accounts for the greatest resistance to<br />

water flux through the root. Plants overcome<br />

this resistance primarily by increasing allocation<br />

to the production <strong>of</strong> new roots (see<br />

Chapter 6).<br />

In dry environments, there is a strong vertical<br />

gradient in soil water potential due to the<br />

low soil water potential in dry surface soils.<br />

However, water moves slowly through the soil<br />

because <strong>of</strong> the low hydraulic conductivity <strong>of</strong> dry<br />

soils. During the day, when plants lose water

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