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Contributions to the debate on water transport¹

⁰ Research School of Biological Sciences, Australian National University, PO Box 475, Canberra 2601, Australia

The useful criticisms of my theory of water transport by Comstock (American Journal of Botany 86: 1077–1081) and by Stiller and Sperry (American Journal of Botany 86: 1082–1086) are acknowledged and reviewed. I make the following responses. (1) Tensile stresses to contain tissue pressure are kept within modest limits by the organization of vascular tissues into cylindrical bundles with small ratios of radius/boundary thickness. (2) The balance of pressures within tissues of a nontranspiring leaf is best understood by treating it as a single compartment containing several pressure-generating engines whose resultant is the pressure throughout the compartment. An error in the published notional balances for a transpiring leaf is corrected. (3) The argument against a valve in the transpiring leaf, which allows water out but not in, is not convincing. (4) The "robust and extremely consistent" cohesion theory gains this status by neglecting large bodies of experimental fact, once well known to plant physiologists. (5) The demonstration that living cells are not involved in the refilling of embolisms in birch stems is welcomed as an important advance. However, the major questions remain unresolved. (6) Proof is still needed that embolisms in vessels are not refilled by the collapse of gas bubbles under small positive pressures during conductance measurement. (7) The survival of unbroken water threads in vessels under centrifugal stress has still not been demonstrated. (8) Both questions 6 and 7 can be easily answered by direct observation of gas/liquid volumes in frozen stems in the cryo-scanning electron microscope.

Key Words: cavitation • cohesion theory • compensating pressure theory • embolism • tissue pressure • water transport

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