Pine Photosynthesis: Needle Longevity, Structural and Physiological Controls, Needle and Tree Age Dependent Changes and Plasticity
摘要
Pines colonize a wide range of habitats from subarctics to tropics, but the comparative information of global variation in pine needle photosynthesis and associated traits has been limited. In this chapter, we first introduce the key functional traits determining needle photosynthesis rate and discuss the ways of expressing needle photosynthesis rates in pines. We use novel global datasets of needle longevity (Λ) and structural, chemical, and physiological traits to assess the overall trait variation across the pines and compare the pine trait variation with global trait variation. We further analyze differences in needle photosynthetic characteristics (photosynthetic capacity per unit area, Aarea, and dry mass, Amass, photosynthetic nitrogen use efficiency, PNUE) in relation to needle structural characteristics (leaf dry mass per unit area, LMA), nitrogen content per dry mass (Nmass) and area (Narea), and Λ, and develop the pine needle economics spectrum. We use the quantitative limitation analysis to characterize the degree to which needle photosynthetic rates are constrained by stomatal and mesophyll diffusion and biochemical limitations, and how the limitations scale with needle functional traits. We conclude by reviewing alterations in needle photosynthetic rates as driven by needle age, plant age and size, and acclimation to environment. The analysis demonstrates about 15-fold variation in key needle traits Aarea, Amass, PNUE, Narea and LMA, with a smaller, 4.5-fold variation in Nmass. Needle average longevity varied almost 30-fold across pines, albeit most pines are characterized by moderate needle longevity of 2–4 yr. Classic leaf economics spectrum relationships were uncovered across pines, characterizing the trade-offs between investments in leaf structural and photosynthetic tissues as driven by variation in needle longevity. In addition to classic economics spectrum traits, available data of different photosynthetic constraints indicated that mesophyll diffusion limitations are greater in pines with more robust needles. Apart from the broad trait relationships, we highlight major impacts of needle ontogeny (juvenile vs adult), age of adult needles, and tree size and environmental acclimation (plasticity) on needle functional traits. Overall, we demonstrate remarkable variation in pine needle photosynthetic capacity across species and during needle and plant lifetime, broadly consistent with their conservative N use strategy and position in the lower end of the leaf economics spectrum.