The Theory of the Rise of Sap in Trees: Some Historical and Conceptual Remarks

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• 作者：Harvey R. Brown
• 关键词：Eugen Askenasy ; Josef B?hm ; Edwin B. Copeland ; Pierre Cruiziat ; Francis Darwin ; Henry H. Dixon ; George Francis FitzGerald ; Stephen Hales ; Taco Hajo van den Honert ; John Joly ; John A. Milburn ; Park S. Nobel ; J.J. Oertli ; William F. Picard ; John S. Sperry ; Ernst Steudle ; Eduard Strasburger ; Melvin T. Tyree ; Martin H. Zimmermann ; cohesion ; tension theory ; rise of sap ; transpiration ; capillarity ; cavitation ; negative xylem pressure ; hydraulic architecture ; global warming ; climate change ; history of
• 刊名：Physics in Perspective (PIP)
• 出版年：2013
• 出版时间：September 2013
• 年：2013
• 卷：15
• 期：3
• 页码：320-358
• 全文大小：444KB
• 参考文献：1. Sir Isaac Newton, / Opticks or A Treatise of the Reflexions, Refractions, Inflexions & Colours of Light. Based on the Fourth Edition London 1730. With a Foreword by Albert Einstein. An Introduction by Sir Edmund Whittaker. A Preface by I. Bernard Cohen (New York: Dover Publications, 1979), Query 31, pp. 375-406, on p. 394.
  2. Francis Darwin, “On the Ascent of Water in Trees,- / Report of the Sixty- / Sixth Meeting of the British Association for the Advancement of Science held at Liverpool in September 1896 (London: John Murray, 1896), 674-683, on 678; reprinted as “Report of a Discussion on the Ascent of Water in Trees,- / Annals of Botany 10 (1896), 630-643, on 635; Vines, “Discussion,-644-647; Joly, “Discussion,-647-660; FitzGerald, “Discussion,-660-661.
  3. J.S. Rowlinson, / Cohesion: A Scientific History of Intermolecular Forces (Cambridge and New York: Cambridge University Press, 2002), p. 262.
  4. Guillermo Angeles, Barbara Bond, John S. Boyer, Tim Brodribb, J. Renée Brooks, Michael J. Burns, Jeannine Cavender-Bares, Mike Clearwater, Hervé Cochard, Jonathan Comstock, Stephen D. Davis, Jean-Christophe Domec, Lisa Donovan, Frank Ewers, Barbara Gartner, Uwe Hacke, Tom Hinckley, N. Michelle Holbrook, Hamlyn G. Jones, Kathleen Kavanagh, Bev Law, Jorge López-Portillo, Claudio Lovisolo, Tim Martin, Jordi Martínez-Vilalta, Stefan Mayr, Frederick C. Meinzer, Peter Melcher, Maurizio Mencuccini, Stephen Mulkey, Andrea Nardini, Howard S. Neufeld, John Passioura, William T. Pockman, R. Brandon Pratt, Serge Rambal, Hanno Richter, Lawren Sack, Sebastiano Salleo, Andrea Schubert, Paul Schulte, Jed P. Sparks, John Sperry, Robert Teskey, and Melvin Tyree, “The Cohesion-Tension Theory,- / New Phytologist 163 (2004), 451-452.
  5. Ulrich Zimmermann, Heike Schneider, Lars H. Wegner, and Axel Haase, “Water ascent in tall trees: does evolution of land plants rely on a highly metastable state?- / New Phytologist 162 (2004), 575-615.
  6. Philippe Gerrienne, Patricia G. Gensel, Christine Strullu-Derrien, Hubert Lardeux, Philippe Steemans, and Cyrille Prestianni, “A Simple Type of Wood in Two Early Devonian Plants,- / Science 333 (2011), 837.
  7. David B. Lindenmayer, William F. Laurance, and Jerry F. Franklin, “Global Decline in Large Old Trees,- / Science 338 (2012), 1305-1306.
  8. George Koch, Stephen Sillett, Gregg Jennings, and Stephen Davis, “How Water Climbs to the Top of a 112 Meter-Tall Tree,-Essay 4.3 (2006), 5 pages, in / Plant Physiology Online, / Fifth Edition; website <http://5e.plantphys.net/article.php?ch=4&id=100>. They note that for California redwoods, summer fog is an important supply of water, and that direct absorption by the leaves may supplement the uptake from root capture of fog drip.
  9. Al Carder, / Forest Giants of the World Past and Present (Markham, Ontario: Fitzhenry & Whiteside, 1995), pp. 1-18, 66-78. Richard McArdle, a former chief of the U.S. Forest Service, estimated a Douglas-fir in Washington to be 120 meters in 1924. The tallest Australian eucalypt today is 100 meters, and there is strong anecdotal evidence of taller ones in the past. The tallest known Sitka spruce ( / Picea sitchensis) in California has a height of nearly 97 meters.
  10. Joel Bourne, “Redwoods. The Super Trees,- / National Geographic 216 (October 2009), 12 pages. A redwood tree’s annual rate of wood production increases with age for at least 1500?years.
  11. T.T. Kozlowski and S.G. Pallardy, / Physiology of Woody Plants (Second Edition San Diego, London, Boston, New York, Sydney, Tokyo, Toronto: Academic Press, 1997), p. 299. It is noteworthy that the oldest trees can live on dry sites; as befits such conditions, bristlecone pines are small, thick trees with highly reduced growth rates.
  12. David Suzuki and Wayne Grady, / Tree: A Life Story (Vancouver, Toronto, Berkeley: Greystone Books, 2004), p. 149.
  13. B.A. Meylan and B.G. Butterfield, / Three- / dimensional structure of wood (Hong Kong: Chapman and Hall, Ltd., 1972), give a spectacular collection of electron microscope photographs of woody structure.
  14. See, for example, Ernst Steudle, “Trees under tension,- / Nature 378 (1995), 663-664, on 663.
  15. Rubin Shmulsky and P. David Jones, / Forest Products and Wood Science: An Introduction. (Sixth Edition New York: John Wiley & Sons, 2011), p. 443, note that the weight of wood used today as a raw material in the United States is still greater than the weight of all metals and plastics combined.
  16. See, for example, Colin Tudge, / The Secret Life of Trees: How They Live and Why They Matter (London: Penguin Press Science, 2006).
  17. Suzuki and Grady, / Tree (ref. 12), p. 68. It is noteworthy that in this elegant and highly readable book on trees, built around the life cycle of a Douglas-fir, the authors present, also in chapter 2, a confused account of the mechanism of sap rise, ultimately regarding it as a “mystery.-/span>
  18. David J Beerling, and Peter J. Franks, “The hidden cost of transpiration,- / Nature 464 (2010), 495-496; Steudle, “Trees under tension-(ref. 14).
  19. Paul J. Kramer and John S. Boyer, / Water Relations of Plants and Soils (San Diego, New York, Boston, London, Sydney, Tokyo, Toronto: Academic Press, 1995), p. 203.
  20. / Ibid., p. 1447; see also Richard A. Betts, “Afforestation cools more or less,- / Nature Geoscience 4 (2011), 504-505.
  21. Historical sketches of varying lengths can be found in Darwin, “Report-and Vines, Joly, and FitzGerald “Discussions-(ref. 2); Edwin Bingham Copeland, “The Rise of the Transpiration Stream: An Historical and Critical Discussion,- / Botanical Gazette 34 (1902), 161-193, 260-283; Henry H. Dixon, / Transpiration and the Ascent of Sap in Plants (London: Macmillan and Co., 1914), Chapter IV, pp. 81-100; Edwin C. Miller, / Plant Physiology; With reference to the green plant (New York and London: McGraw-Hill Book Company, 1938), pp. 855-872; Kozlowski and Pallardy, / Physiology of Woody Plants (ref. 11), pp. 259-260; K.N.H. Greenidge, “Ascent of sap,- / Annual Review of Plant Physiology 8 (1957), 237-256; William F. Pickard, “The Ascent of Sap in Plants,- / Progress in Biophysics and Molecular Biology 37 (1981), 181-229; and especially the online essay by Hanno Richter and Pierre Cruiziat, “A Brief History of the Study of Water Movement in the Xylem,-Essay 4.1 (2002), in / Plant Physiology Online, Fifth Edition, 5 pages, website <http://5e.plantphys.net/article.php?ch=&id=98>.
  22. Succinct outlines of the CT theory are found in N. Michele Holbrook, Maclej A. Zwieniecki, and Peter J. Melcher, “The Dynamics of ‘Dead Wood- Maintenance of Water Transport Through Plant Stems,- / Integrative and Comparative Biology 42 (2002), 492-496, and Pierre Cruiziat and Hanno Richter, “The Cohesion-Tension Theory at Work,-Essay 4.2 (2006), in / Plant Physiology Online, Fifth Edition, 4 pages, website <http://5e.plantphys.net/article.php?ch=4&id=99>. More detailed review papers are Greenidge, “Ascent of sap-(ref. 21); M.J. Canny, “Flow and Transport in Plants,- / Annual RevXylemiew of Fluid Mechanics 9 (1977), 275-296; and Ernst Steudle, “The Cohesion-Tension Mechanism and the Acquisition of Water by Plant Roots,- / Annual Review of Plant Physiology and Plant Molecular Biology 52 (2001), 847-875. Lengthier treatments are found in in the textbooks of Kramer and Boyer, / Water Relations (ref. 19), pp. 234-251, Kozlowski and Pallardy, / Physiology of Woody Plants (ref. 11), pp. 259-260; M.T. Tyree and M.H. Zimmermann, / Xylem Structure and the Ascent of Sap (Berlin, Heidelberg, New York: Springer-Verlag, 2002), pp. 49-88, this being the second edition of the authoritative Martin H. Zimmermann, / Xylem Structure and the Ascent of Sap (Berlin, Heidlberg, New York, Tokyo: Springer Verlag, 1983), pp. 37-65; Wilfried Ehlers and Michael Goss, / Water Dynamics in Plant Production (Wallingford and Cambridge, Mass.: CABI Publishing, 2003), pp. 82-84; and Park S. Nobel, / Physicochemical and Environmental Plant Physiology (Third Edition Amsterdam: Elsevier Academic Press, 2005), p. 459. For a physicist, the technical review paper by Pickard, “Ascent of Sap in Plants-(ref. 21), may be the most informative and satisfying, though it is somewhat out of date, particularly in relation to mechanisms for refilling embolized xylem conduits. An excellent, up-to-date, somewhat less technical review is John S. Sperry, “Hydraulics of Vascular Water Transport,-in Przemyslaw Wojtaszek, ed., / Mechanical Integration of Plant Cells and Plants [ / Signaling and Communication in Plants, Vol. 9] (Berlin and Heidelberg: Springer-Verlag, 2011), pp. 303-327. The literature is huge; these publications represent a very small selection; a useful list of key papers is found in Angeles, / et al., “Cohesion-Tension Theory-(ref. 4).
  23. Quoted in Ehlers and Goss, / Water Dynamics (ref. 22), p. 82, who attribute this quotation to Josef B?hm, “Capillarit?t und Saftsteigen,- / Bericht der Deutschen Botanischen Gesellschaft 11 (1893), 203-212, on 207.
  24. Eduard Strasburger, / Ueber den Bau und die Verrichtungen der Leitungsbahnen in den Pflanzen (Jena: Gustav Fischer, 1891); / idem, / Ueber das Saftsteigen (Jena: Gustav Fischer, 1893).
  25. Darwin, “Report-(ref. 2), p. 640.
  26. / Ibid., p. 631; Copeland, “Rise of the Transpiration Stream-(ref. 21), provides much detail on the range of ideas and experimental evidence being brought to bear on the issue in the 19th century -and his own skepticism about the validity of the CT theory.
  27. Herman Helmholtz, “Ueber galvanische Polarisation in gasfreien Flüssigkeiten-[1873], in / Wissenschaftliche Abhandlungern. Erster Band (Leipzig: Johann Ambrosius Barth, 1882), pp. 821-834, on p. 830.
  28. B?hm, “Capillarit?t und Saftsteigen,-(ref. 23), p. 206. For a useful summary of B?hm’s energetic experimental work on the rise of sap, see Ehlers and Goss, / Water Dynamics (ref. 22), pp. 82-84. The translation from the German of these passages is due to John Pannell (private communication).
  29. Patrick N. Wyse Jackson, “A Man of Invention: John Joly (1857-1933), Engineer, Physicist and Geologist,-in David Scott, ed., / Treasures of the Mind. A Trinity College Dublin Quatercentenary Exhibition (London: Sotheby’s, 1992), pp. 89-96.
  30. Michael Jones, “‘A Name Writ in Water- Henry Horatio Dixon 1869-1953,-in Scott, / Treasures of the Mind (ref. 29), pp. 97-103; online at website <dixon.php" class="a-plus-plus">http://www.tcd.ie/Botany/tercentenary/300-years/chairs/henry-horatio-dixon.php>, 5 pages.
  31. Henry H. Dixon and J. Joly, “On the Ascent of Sap-(Abstract), / Proceedings of the Royal Society of London 57 (1894), 3-5.
  32. Henry H. Dixon and John Joly, “On the Ascent of Sap,- / Philosophical Transactions of the Royal Society of London B 186 (1895), 563-576.
  33. / Ibid., p. 563.
  34. Copeland, “Rise of the Transpiration Stream-(ref. 21), p. 186. E. Askenasy, “Ueber das Saftsteigen,- / Verhandlungen des Naturhistorisch- / Medizinischen Vereins zu Heidelberg 5 (1896), 325-345; / idem, “Beitrage zur Erklrung des Saftsteigens,- / ibid., 429-448.
  35. Joly, “Discussion-(ref. 2), p. 647.
  36. For details, see Darwin, “Report-(ref. 2), pp. 635-642; Vines, “Discussion: (ref. 2), pp. 646-647; Joly, “Discussion-(ref. 2), pp. 647-654; FitzGerald, “Duscussion: (ref. 2), pp. 660-661; see also Pickard, “Ascent of Sap in Plants-(ref. 21), p. 220.
  37. Pickard, “Ascent of Sap in Plants-(ref. 21), p. 185.
  38. Joly, “Discussion-(ref. 2), p. 648.
  39. Richter and Cruiziat, “Brief History-(ref. 21), p. 2 of 5.
  40. / Ibid.
  41. Darwin, “Report-(ref. 2), p. 635.
  42. Joly, “Discussion-(ref. 2), pp. 648-649.
  43. See, for example, Steudle, “Cohesion-Tension Mechanism-(ref. 22), p. 854.
  44. Joly, “Discussion-ref. 2), p. 652, where Joly refers to a model involving a porous pot connected to a supply of water, but the argument is supposed to illuminate what happens in trees.
  45. Pickard, “Ascent of Sap in Plants-(ref. 21), pp. 220-221.
  46. Joly, “Discussion-(ref. 2), p. 659.
  47. Dixon, / Transpiration (ref. 21), p. 29. It is noteworthy that the five B?hm papers cited by Dixon do not include his 1893 paper.
  48. Stephen Hales, / Vegetable Staticks: Or, An Account of some Statical Experiments on the Sap in VEGETABLES, etc. (London: W. and J. Innys and T. Woodward, 1727; reprinted London: Macdonald and New York: American Elsevier, 1969).
  49. R. Harré, / The Method of Science. A Course in Understanding Science, based upon the De Magnete / of William Gilbert, and the Vegetable Staticks / of Stephen Hales (London and Winchester: Wykeham Publications and New York: Springer-Verlag, 1970), p. 65. He includes a short biography of Hales in Chapter 7, pp. 70-76; amongst Hales’s many contributions to science was the discovery of carbon dioxide in air.
  50. Franz Floto, “Stephen Hales and the cohesion theory,- / Trends in Plant Science 4 (1999), 209.
  51. Hales, / Vegetable Staticks (ref. 48), p. 43.
  52. / Ibid., p. 56. It seems that Hales himself may not have been the first to connect capillarity with the rise of sap; see Dixon, / Transpiration (ref. 21), p. 27, who refers to the 1723 views of Christian Wolff who “believed that the forces involved were the expansion of air and capillarity.-
  53. Rowlinson, / Cohesion (ref. 3), p. 264.
  54. / Ibid., Section?3.2, pp. 86-102, for an account of their work, particularly related to capillarity.
  55. For details of the early work in the field, see George S. Kell, “Early observations of negative pressures in liquids,- / American Journal of Physics 51 (1983), 1038-1041.
  56. David H. Trevena, “Marcelin Berthelot’s First Publication in 1850, on the Subjection of Liquids to Tension,- / Annals of Science 35 (1978), 45-54.
  57. Hales, / Vegetable Staticks (ref. 48), p. 9; Darwin, “Report-(ref. 2), p. 630.
  58. See, in particular. Steudle, “Trees under tension-(ref. 14), p. 663, who suggests that nonbotanists typically make the mistake of attributing sap rise to, if anything, capillary action in the xylem.
  59. Hales, / Vegetable Staticks (ref. 48), pp. 77-78. It is hard to reconcile this passage with the claim by Beerling and Franks, “The hidden cost of transpiration-(ref. 18), p. 496, that Hales, despite being aware of the role of “perspiration-in the ascent of sap, did not discover that plants transpire water from leaves.
  60. P.F. Scholander, Warner E. Love, and John W. Kanwisher, “The Rise of Sap in Tall Grapevines,- / Plant Physiology 30 (1955), 93-104. Studies include that of the common northern grapevine ( / Vitis labrusca), which can sometimes attain heights of 17 to 18 meters in wooded areas in Massachusetts.
  61. As suggested by William F. Pickard, “How might a Tracheary Element which is Embolized by Day be Healed by Night?- / Journal of Theoretical Biology 141 (1989), 259-279; the positive-pressure mechanism in tall vines is based on temperature-associated osmotic water uptake from rehydrated cells of the bark. It is noteworthy that Hales, / Vegetable Staticks (ref. 48), Chapter III, pp. 57-68, performed the first published experiments designed to measure root pressure, in grapevines ( / Vitis vinifera).
  62. Pickard, “How might a Tracheary Element-(ref. 61), p. 264, and Melvin T. Tyree; Sebastiano Salleo; Andrea Nardini; Maria Assunta Lo Gullo, and Roberto Mosca, “Refilling of Embolized Vessels in Young Stems of Laurel. Do We Need a New Paradigm?- / Plant Physiology 120 (1999), 11-21.
  63. Nobel, / Physicochemical and Environmental Plant Physiology (ref. 22), p. 53.
  64. Pickard, “How might a Tracheary Element-(ref. 61), p. 268.
  65. S.H. Vines, “The Suction-force of Transpiring Branches,- / Annals of Botany 10 (1896), 429-444. The happily-named Vines was measuring significant tension within leafless branches- see Vines, “Discussion: (ref. 2), pp. 644-647. This capillarity view has a certain amount in common with the “imbibition-theory of sap rise that was prominent prior to the 1880s and largely associated with the name of the great German botanist Julius von Sachs (1832-1897), though this theory held that the flow of sap takes place within the walls of xylem conduits; see Copeland, “Rise of the Transpiration Stream-(ref. 21), pp. 179-181, and Dixon / Transpiration, (ref. 21), pp. 83-84.
  66. For remarks on the farsighted 1886 work of N?geli, see Copeland, “Rise of the Transpiration Stream-(ref. 21), pp. 183-185.
  67. See in particular the survey of results in Greenidge “Ascent of sap-(ref. 21), pp. 238-249.
  68. Tyree and Zimmermann, / Xylem Structure (ref. 22), p. 62.
  69. M.F. Donny, “On the Cohesion of Liquids and their Adhesion to Solid Bodies,- / Philosophical Magazine 28 (1846), [291]-294. Hasok Chang, / Inventing Temperature: Measurement and Scientific Progress (Oxford and New York: Oxford University Press, 2004), pp. 8-39, provides a fascinating treatment of the history of the theory of boiling.
  70. J.J. Oertli, “The stability of water under tension in the xylem,- / Zeitschrift für Pflanzenphysiologie 65 (1971), 195-209, on 208.
  71. / Ibid., p. 196, and Pickard, “Ascent of Sap in Plants-(ref. 21), Section IV, pp. 198-208.
  72. Oertli, “The stability of water-(ref. 70), p. 207. The point seems to be widely accepted. “The conditions for heterogeneous nucleation at preexistent sites appear to be equally hard to fulfill: the tracheary element is presumably water filled from earliest differentiation with the result that its putatively hydrophilic boundaries should be thoroughly wet and unlikely to stabilize nuclei-- see Pickard, “Ascent of Sap in Plants-(ref. 21), pp. 205, 195. “Xylem conduits are water filled from inception and contain no entrapped air bubbles that could nucleate cavitation- see M.T. Tyree and J.S. Sperry, “Vulnerability of Xylem to Cavitation and Embolism,- / Annual Review of Plant Physiology and Plant Molecular Biology 40 (1989), 19-38, on 20.
  73. Kramer and Boyer, / Water Relations (ref. 19), p. 203. It should be noted that this remark occurs in the context of a discussion of how important transpiration is for the growth of a plant.
  74. Sperry, “Hydraulics-(ref. 22), p. 308.
  75. Steudle, “Cohesion-Tension Mechanism-(ref. 22), p. 854.
  76. For a review of the mechanism of cell growth in plants, and the role of osmosis therein, see J.S. Boyer, “Water Transport,- / Annual Review of Plant Physiology 36 (1985), 473-516, on 492-500, and John S. Boyer, A.J. Cavalieri, and E.-D. Schulze, “Control of the rate of cell enlargement: Excision, wall relaxation, and growth-induced water potentials,- / Planta 163 (1985), 527-543. John Sperry has pointed out to me (private communication) that in cell growth, as opposed to transpiration, the air-water meniscus is stressed from below, but in both cases the pressure drop is most proximally generated by the air-water meniscus by way of capillary action.
  77. J.S. Boyer, “Cell enlargement and growth-induced water potentials,- / Physiologia Plantarum 73 (1988), 311-316.
  78. Dixon and Joly, “On the Ascent of Sap-(ref. 32), p. 568.
  79. These results were reiterated in Dixon, / Transpiration (ref. 21), p. 87.
  80. Dixon and Joly, “On the Ascent of sap-(ref. 32), p. 572.
  81. Tyree and Sperry, “Vulnerability of Xylem-(ref. 72), p. 26. Apart from vines, root pressure can act to recover embolized xylem conduits in herbs, shrubs, and small trees; it has never been recorded in gymnosperms or in most forest trees; see Tyree, Salleo, Nardini, Assunta Lo Gullo, and Mosca, “Refilling of Embolized Vessels-(ref. 62).
  82. Dixon, / Transpiration (ref. 21), p. 210.
  83. Pickard, “Ascent of Sap in Plants-(ref. 21), pp. 205-208.
  84. Tyree and Sperry, “Vulnerability of Xylem-(ref. 72).
  85. For further details on the nature of cavitation, see Tyree and Zimmermann, / Xylem Structure (ref. 22), section?4.1, pp. 89-94.
  86. Hales, / Vegetable Staticks (ref. 48), Chapter IV, pp. 69-84.
  87. Pierre Cruiziat, Hervé Cochard, and Thierry Améglio, “Hydraulic architecture of trees: main concepts and results,- / Annals of Forrest Science 59 (2002), 723-752, on 736.
  88. For a brief review of this mechanism, see James K. Wheeler and N. Michele Holbrook, “Cavitation and Refilling,-Essay 4.4 (2007), in / Plant Physiology Online, Fifth Edition, 6 pages, website <http://5e.plantphys.net/article.php?ch=4&id=395>; for more detail see also Cruiziat, Cochard, and Améglio, “Hydraulic architecture of trees-(ref. 87), pp. 734-741, and especially Holbrook, Zwieniecki, and Melcher, “Dynamics-(ref. 22), pp. 493-495.
  89. Clark L. Stevens and Russell L. Eggert, “Observations on the Causes of the Fow of Sap in Red Maple,- / Plant Physiology 20 (1945), 636-648, on 647-648.
  90. For a useful review, see Tyree and Zimmermann, / Xylem Structure (ref. 22), Section?3.9, pp. 81-88.
  91. For details of the two main competing models, and recent anatomical evidence in favor of the 1995 osmotic model due to Tyree, see Damien Cirelli; Richard Jagels, and Melvin T. Tyree, “Toward an improved model of maple sap exudation: the location and role of osmotic barriers in sugar maple, butternut and white birch,- / Tree Physiology 28 (2008), 1145-1155.
  92. Pickard, “Ascent of Sap in Plants-(ref. 21), p. 223.
  93. John A. Milburn, “Sap Ascent in Vascular Plants: Challengers to the Cohesion Theory Ignore the Significance of Immature Xylem and the Recycling of Münch Water,- / Annals of Botany 78 (1996), 399-407.
  94. U. Zimmermann, F.C Meinzer, R. Bankert, J.J. Zhu, H. Schneider, G. Goldstein, E. Kuchenbrod, and A. Haase, “Xylem water transport: is the available evidence consistent with the cohesion theory?- / Plant, Cell and Environment 17 (1994), 1169-1181, and Ulrich Zimmermann, Frederick. Meinzer, and Freidrich-Wilhelm Bentrup, “How Does Water Ascend in Tall Trees and Other Vascular Plants?- / Annals of Botany 76 (1995), 545-551.
  95. John S. Sperry, “Limitations on Stem Water Transport and Their Consequences,-in Barbara L. Gartner, ed., / Plant Stems: Physiology and Functional Morphology (San Diego, New York, Boston, London, Sydney, Tokyo, Toronto: Academic Press, 1995), pp. 105-124, and J.S. Sperry, N.Z. Saliendra, W.T. Pockman, H. Cochard, P. Cruiziat, S.D. Davis, F.W. Ewers, and M.T. Tyree, “New evidence for large negative xylem pressures and their measurement by the pressure chamber method,- / Plant, Cell and Environment 19 (1996), 427-436.
  96. William T. Pockman, John S. Sperry, and James W. O’Leary, “Sustained and significant negative water pressure in xylem,- / Nature 378 (1995), 715-716.
  97. Milburn, “Sap Ascent-(ref. 93); Sperry, Saliendra, Pockman, Cochard, Cruiziat, Davis, Ewers, and Tyree, “New evidence-(ref. 95), pp. 432-435; Melvin T. Tyree, The Cohesion-Tension theory of sap ascent: current controversies,- / Journal of Experimental Botany 48 (1997), 1753-1765; Chunfang Wei, Ernst Steudle, and Melvin T. Tyree, “Water ascent in plants: do ongoing controversies have a sound basis?- / Trends in Plant Science 4 (1999), 372-375, and the reply to the prior paper in Ulrich Zimmermann, Hans-Jürgen Wagner, Heike Schneider, Markus Rokitta, Axel Haase, and Freidrich-Wilhelm Bentrup, “Water ascent in plants: the ongoing debate,- / ibid. 5 (2000), 145-146; for a detailed analysis of the pressure-probe tool, see A. Deri Tomos and Roger A. Leigh, “THE PRESSURE PROBE: A Versatile Tool in Plant Cell Physiology,- / Annual Review of Plant Physiology and Plant Molecular Biology 50 (1999), 447-472.
  98. See Harvey R. Brown, / Physical Relativity: Space- / time structure from a Dynamical Perspective (Oxford: Clarendon Press, 2005), pp. 86-87, for the similar episode in the history of the special theory of relativity involving Kauffmann’s experimental “refutation-of the Lorentz-Einstein theory in experiments performed between 1901 and 1905.
  99. T.H. van den Honert, “Water Transport in Plants as a Catenary Process,-Discussions of the Faraday Society 3 (1948), 146-153.
  100. A review of the hydraulics of leaves, which represent an important hydraulic bottleneck in trees, is found in Lawren Sack and N. Michele Holbrook, “Leaf Hydraulics,- / Annual Review of Plant Biology 57 (2006), 361-381, and Athena D. McKown; Hervé Cochard, and Lawren Sack, “Decoding Leaf Hydraulics with a Spatially Explicit Model: Principles of Venation Architecture and Implications for Its Evolution,- / The American Naturalist 175 (2010), 447-460. The latter is the first detailed examination of the hydraulic consequences and implications of key leaf venation traits for the economics, ecology, and evolution of plant transport capacity.
  101. Holbrook, Zwieniecki, and Melcher, “Dynamics-(ref. 22), p. 495, and further references therein.
  102. For reviews, see Melvin T. Tyree and Frank W. Ewers, “The hydraulic architecture of trees and other woody plants,- / New Phytologist 119 (1991), 345-360, and Cruiziat, Cochard, and Améglio, “Hydraulic architecture of trees-(ref. 87).
  103. Frederick C. Meinzer; Shelley A. James, and Guillermo Goldstein, “Dynamics of transpiration, sap flow and use of stored water in tropical forest canopy trees,- / Tree Physiology 24 (2004), 901-909.
  104. F.C. Meinzer, J.R. Brooks, J.-C. Domec, B.L. Gartner, J.M. Warren, D.R. Woodruff, K. Bible, and D.C. Shaw, “Dynamics of water transport and storage in conifers studied with deuterium and heat tracing techniques,- / Plant, Cell and Environment 29 (2006), 105-114, on 105.
  105. For more details, see Tyree and Zimmermann, / Xylem Structure (ref. 22), Section?4.8, pp. 132-141. For a review of water storage in plants, see N. Michele Holbrook, “Stem Water Storage,-in Gartner, / Plant Stems (ref. 95), pp. 151-174.
  
• 作者单位：Harvey R. Brown (1)
  
  1. Faculty of Philosophy, University of Oxford, Radcliffe Humanities, Radcliffe Observatory Quarter, Woodstock Road, Oxford, OX2 6GG, UK
  
• ISSN：1422-6960

文摘

The ability of trees to suck water from roots to leaves, sometimes to heights of over a hundred meters, is remarkable given the absence of any mechanical pump. In this study I deal with a number of issues, of both a historical and conceptual nature, in the orthodox Cohesion-Tension (CT) theory of the ascent of sap in trees. The theory relies chiefly on the exceptional cohesive and adhesive properties of water, the structural properties of trees, and the role of evaporation (“transpiration- from leaves. But it is not the whole story. Plant scientists have been aware since the inception of the theory in the late 19th century that further processes are at work in order to “prime-the trees, the main such process -growth itself -being so obvious to them that it is often omitted from the story. Other factors depend largely on the type of tree, and are not always fully understood. For physicists, in particular, it may be helpful to see the fuller picture, which is what I attempt to provide in nontechnical terms.