, Volume 177, Issue 2, pp 321–331 | Cite as

Growth maximization trumps maintenance of leaf conductance in the tallest angiosperm

  • George W. Koch
  • Stephen C. Sillett
  • Marie E. Antoine
  • Cameron B. Williams
Physiological ecology - Original research


Structural and physiological changes that occur as trees grow taller are associated with increased hydraulic constraints on leaf gas exchange, yet it is unclear if leaf-level constraints influence whole-tree growth as trees approach their maximum size. We examined variation in leaf physiology, leaf area to sapwood area ratio (L/S), and annual aboveground growth across a range of tree heights in Eucalyptus regnans. Leaf photosynthetic capacity did not differ among upper crown leaves of individuals 61.1–92.4 m tall. Maximum daily and integrated diurnal stomatal conductance (g s) averaged 36 and 34 % higher, respectively, in upper crown leaves of ~60-m-tall, 80-year-old trees than in ~90-m-tall, 300-year-old trees, with larger differences observed on days with a high vapor pressure deficit (VPD). Greater stomatal regulation in taller trees resulted in similar minimum daily leaf water potentials (Ψ L) in shorter and taller trees over a broad range of VPDs. The long-term stomatal limitation on photosynthesis, as inferred from leaf δ 13C composition, was also greater in taller trees. The δ 13C of wood indicated that the bulk of photosynthesis used to fuel wood production in the main trunk and branches occurred in the upper crown. L/S increased with tree height, especially after accounting for size-independent variation in crown structure across 27 trees up to 99.8 m tall. Despite greater stomatal limitation of leaf photosynthesis in taller trees, total L explained 95 % of the variation in annual aboveground biomass growth among 15 trees measured for annual biomass growth increment in 2006. Our results support a theoretical model proposing that, in the face of increasing hydraulic constraints with height, whole-tree growth is maximized by a resource trade-off that increases L to maximize light capture rather than by reducing L/S to sustain g s.


Eucalyptus regnans Growth maximization Hydraulic balance Leaf area to sapwood area ratio Tree height 



This research was supported by grants from the National Science Foundation (IOS-0445277 and IOS-1010769) and the endowment creating the Kenneth L. Fisher Chair in Redwood Forest Ecology at Humboldt State University. Comments by Mike Ryan and two anonymous reviewers improved the manuscript. We thank Tom Greenwood, Giacomo Renzullo, Jim Campbell-Spickler, Joe Harris, and Robert Van Pelt for help with tree climbing and sampling, and Jason Beringer and Darren Hocking of Monash University for providing micrometeorological data from the Wallaby Creek eddy-covariance tower. Russell D. Kramer helped with statistical analysis using R. We are especially grateful to Ion Maher and Tony Fitzgerald of Kinglake National Park, Victoria, for research permission, hospitality, and logistical assistance.


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Copyright information

© Springer-Verlag Berlin Heidelberg 2014

Authors and Affiliations

  • George W. Koch
    • 1
  • Stephen C. Sillett
    • 2
  • Marie E. Antoine
    • 3
  • Cameron B. Williams
    • 4
  1. 1.Center for Ecosystem Science and SocietyNorthern Arizona UniversityFlagstaffUSA
  2. 2.Department of Forestry and Wildland ResourcesHumboldt State UniversityArcataUSA
  3. 3.Department of Biological SciencesHumboldt State UniversityArcataUSA
  4. 4.Department of Integrative BiologyUniversity of CaliforniaBerkeleyUSA

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