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How strong is intracanopy leaf plasticity in temperate deciduous trees?¹

²Department of Botany, University of Hawaii at Manoa, 3190 Maile Way, Honolulu, Hawaii 96822 USA; ³Arnold Arboretum of Harvard University, 125 Arborway, Jamaica Plain, Massachusetts 02130 USA; ⁴Biology Department, Ithaca College, Center for Natural Sciences, Ithaca, New York 14850 USA; ⁵Department of Organismic and Evolutionary Biology, Harvard University, Biological Laboratories, 16 Divinity Avenue, Cambridge, Massachusetts 02138 USA; ⁶Department of Environmental Science, Policy, and Management, University of California, Berkeley, California 94720 USA

ABSTRACT

Intracanopy plasticity in tree leaf form is a major determinant of whole-plant function and potentially of forest understory ecology. However, there exists little systematic information for the full extent of intracanopy plasticity, whether it is linked with height and exposure, or its variation across species. For arboretum-grown trees of six temperate deciduous species averaging 13–18 m in height, we quantified intracanopy plasticity for 11 leaf traits across three canopy locations (basal-interior, basal-exterior, and top). Plasticity was pronounced across the canopy, and maximum likelihood analyses indicated that plasticity was primarily linked with irradiance, regardless of height. Intracanopy plasticity (the quotient of values for top and basal-interior leaves) was often similar across species and statistically indistinguishable across species for several key traits. At canopy tops, the area of individual leaves was on average 0.5–0.6 times that at basal-interior, stomatal density 1.1–1.5 times higher, sapwood cross-sectional area up to 1.7 times higher, and leaf mass per area 1.5–2.2 times higher; guard cell and stomatal pore lengths were invariant across the canopy. Species differed in intracanopy plasticity for the mass of individual leaves, leaf margin dissection, ratio of leaf to sapwood areas, and stomatal pore area per leaf area; plasticity quotients ranged only up to 2. Across the six species, trait plasticities were uncorrelated and independent of the magnitude of the canopy gradient in irradiance or height and of the species' light requirements for regeneration. This convergence across species indicates general optimization or constraints in development, resulting in a bounded plasticity that improves canopy performance.

Key Words: guard cells • hydraulic limitations • leaf size • leaf shape • plant intelligence • shade tolerance • stomata • succession

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