A hydraulic–photosynthetic model based on extended HLH and its application to Coast redwood (Sequoia sempervirens)

Although recent investigations [Ryan, M.G., Yoder, B.J., 1997. Hydraulic limits to tree height and tree growth. Bioscience 47, 235–242; Koch, G.W., Sillett, S.C.,Jennings, G.M.,Davis, S.D., 2004. The limits to tree height. Nature 428, 851–854; Niklas, K.J., Spatz, H., 2004. Growth and hydraulic (not mechanical) constraints govern the scaling of tree height and mass. Proc. Natl Acad. Sci. 101, 15661–15663; Ryan, M.G., Phillips, N., Bond, B.J., 2006. Hydraulic limitation hypothesis revisited. Plant Cell Environ. 29, 367–381; Niklas, K.J., 2007. Maximum plant height and the biophysical factors that limit it. Tree Physiol. 27, 433–440; Burgess, S.S.O., Dawson, T.E., 2007. Predicting the limits to tree height using statistical regressions of leaf traits. New Phytol. 174, 626–636] suggested that the hydraulic limitation hypothesis (HLH) is the most plausible theory to explain the biophysical limits to maximum tree height and the decline in tree growth rate with age, the analysis is largely qualitative or based on statistical regression. Here we present an integrated biophysical model based on the principle that trees develop physiological compensations (e.g. the declined leaf water potential and the tapering of conduits with heights [West, G.B., Brown, J.H., Enquist, B.J., 1999. A general model for the structure and allometry of plant vascular systems. Nature 400, 664–667]) to resist the increasing water stress with height, the classical HLH and the biochemical limitations on photosynthesis [von Caemmerer, S., 2000. Biochemical Models of Leaf Photosynthesis. CSIRO Publishing, Australia]. The model has been applied to the tallest trees in the world (viz. Coast redwood (Sequoia sempervirens)). Xylem water potential, leaf carbon isotope composition, leaf mass to area ratio at different heights derived from the model show good agreements with the experimental measurements of Koch et al. [2004. The limits to tree height. Nature 428, 851–854]. The model also well explains the universal trend of declining growth rate with age.