The necessity to protect the xylem transport system against theeffects of cavitation and airlocks may explain why tracheids havepersisted in the flowering plants alongside vessels. The vessels, withno barriers to water flow between cell and cell, offer much lessresistance to water flow than tracheids, where water must passfrom tracheid to tracheid through the cell wall, mainly throughpits. Moreover some vessel elements are extremely narrow, therebeing normally a range of widths of vessel elements in the sameplant organ. Volume flow through a tube is proportional to the fourthpower of the radius of the tube. Hence a two-fold increase in the radiusof a vessel from say, 5 mm to 10 mm would increase volume flow (forthe same tension) 16-fold (24 = 16). Since wide vessels are so muchmore efficient for a bulk flow of water, why should a vascular bundlecontain, in addition to wide vessels, narrow vessels and tracheids?The most probable answer is that this gives the plant the flexibility toreact efficiently to varying environmental water status. When thesoil C is high the plant does not require very high tensions in thexylem to extract the water; in such a situation, most of the transpir-ation stream would pass through the widest vessels, which offer the least resistance. But when water stress sets in and xylem tensionincreases, it will be the widest vessels that are the most vulnerableto cavitation. The narrower vessels and the tracheids then can takeover the function of water conduction; the same high tensions whichcause vessels to cavitate also overcome the resistance in the narrowerconducting cells. In tracheids, any cavitation event is confined to onesingle cell, and tracheids are also relatively narrow, so they are theleast vulnerable to water stress.The movement of cohesive water columns in the xylem undertranspiration pull may thus be buffered against serious disruptionfrom cavitation by excess capacity; by the regular annual replace-ment of old xylem by new in perennials; by refilling of air-filledvessels by root pressures, by capillarity and by pressure from adjacentliving cells; by the bypassing of airlocks in cell walls; and by thepresence in xylem of tracheids and narrow vessels, which are lesssusceptible to cavitation. There is not enough evidence for acceptingalternative theories for the ascent of sap which deny the existence ofhigh tensions in the xylem. Nevertheless, data from the pressure-probe measurements, and other apparently anomalous observations,are drawing attention to the possible functions of the living cells ofthe xylem, and of the neighbouring phloem, in water movement.
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