Transpiration-Pull
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In 1895, the Irish plant physiologists H. H. Dixon and J. Joly proposed that water is pulled up the plant by tension (negative pressure) from above.
As we have seen, water is continually being lost from leaves by transpiration. Dixon and Joly believed that the loss of water in the leaves exerts a pull on the water in the xylem ducts and draws more water into the leaf.
But even the best vacuum pump can pull water up to a height of only 34 ft or so. This is because a column of water that high exerts a pressure (~15 lb/in2) just counterbalanced by the pressure of the atmosphere. How can water be drawn to the top of a sequoia (the tallest is 370 feet high)? Taking all factors into account, a pull of at least 270 lb/in2 is probably needed.
The answer to the dilemma lies the cohesion of water molecules; that is the property of water molecules to cling to each through the hydrogen bonds they form.
When water is confined to tubes of very small bore, the force of cohesion between water molecules imparts great strength to the column of water. Tensions as great as 3000 lb/in2 are needed to break the column, about the value needed to break steel wires of the same diameter. In a sense, the cohesion of water molecules gives them the physical properties of solid wires.
Because of the critical role of cohesion, the transpiration-pull theory is also called the cohesion theory.
Some support for the theory
* If sap in the xylem is under tension, we would expect the column to snap apart if air is introduced into the xylem vessel by puncturing it. This is the case.
* If the water in all the xylem ducts is under tension, there should be a resulting inward pull (because of adhesion) on the walls of the ducts. This inward pull in the band of sapwood in an actively transpiring tree should, in turn, cause a decrease in the diameter of the trunk.
* The rattan vine may climb as high as 150 ft on the trees of the tropical rain forest in northeastern Australia to get its foliage into the sun. When the base of a vine is severed while immersed in a basin of water, water continues to be taken up. A vine less than 1 inch in diameter will "drink" water indefinitely at a rate of up to 12 ml/minute.
If forced to take water from a sealed container, the vine does so without any decrease in rate, even though the resulting vacuum becomes so great that the remaining water begins to boil spontaneously. (The boiling temperature of water decreases as the air pressure over the water decreases, which is why it takes longer to boil an egg in Denver than in New Orleans.)
* Transpiration-pull enables some trees and shrubs to live in seawater. Seawater is markedly hypertonic to the cytoplasm in the roots of the coastal mangrove, and we might expect water to leave the cells resulting in a loss in turgor and wilting. In fact, the remarkably high tensions (on the order of 500–800 lb/in2) in the xylem can pull water into the plant against this osmotic gradient. So mangroves literally desalt seawater to meet their needs.
Problems with the theory
When water is placed under a high vacuum, any dissolved gases come out of solution as bubbles (as we saw above with the rattan vine). This is called cavitation. Any impurities in the water enhance the process. So measurements showing the high tensile strength of water in capillaries require water of high purity — not the case for sap in the xylem.
So might cavitation break the column of water in the xylem and thus interrupt its flow? Probably not so long as the tension does not greatly exceed 270 lb/in2.
By spinning branches in a centrifuge, it has been shown that water in the xylem avoids cavitation at negative pressures exceeding 225 lb/in2.
Sequoias can successfully lift water 358 ft (109 m) — which would require a tension of 270 lb/in2.
However, such heights may be approaching the limit for xylem transport. (The tallest tree ever measured, a Douglas fir, was 413 ft. high.) Measurements close to the top of the tallest living sequoia (370 ft high) show that the high tensions needed to get water up there have resulted in:
* smaller stomatal openings, causing
* lower concentrations of CO2 in the needles, causing
* reduced photosynthesis, causing
* reduced growth (smaller cells and much smaller needles).