Spatial and age-dependent modifications of photosynthetic capacity in four Mediterranean oak species
Ülo Niinemets A B , John D. Tenhunen C and Wolfram Beyschlag DA Department of Plant Physiology, University of Tartu, Riia 23, Tartu 51 010, Estonia.
B Centro di Ecologia Alpina, I-38 040 Viote del Monte Bondone (TN), Italy. Corresponding author. Email: ulo.niinemets@ut.ee
C Department of Plant Ecology, University of Bayreuth, D-95 440, Bayreuth, Germany.
D Department of Experimental and Systems Ecology, University of Bielefeld, Universitätsstraße 25, 33 615 Bielefeld, Germany.
Functional Plant Biology 31(12) 1179-1193 https://doi.org/10.1071/FP04128
Submitted: 15 July 2004 Accepted: 21 September 2004 Published: 8 December 2004
Abstract
Drought is one of the most important limitations of photosynthesis in Mediterranean climates. However, Mediterranean sclerophyllous species with long-lived leaves also support extensive and dynamic canopies, with potentially large spatial and age-dependent gradients. We studied within-canopy and temporal patterns in foliage structure, chemistry and photosynthesis in the evergreen species Quercus coccifera L., Q. ilex L. subsp. ballota (Desf.) Samp. in Bol. and Q. suber L. and in the semi-deciduous marcescent species Q. faginea Lam. to determine the role of within-canopy shading and leaf age on foliage functioning. There was a 2.5-fold within-canopy gradient in leaf dry mass per unit area (MA) that was accompanied by a 3-fold range in area-based leaf nitrogen (N) content, the capacity for photosynthetic electron transport (Jmax) and maximum Rubisco carboxylase activity (Vcmax), while the fractional investments of leaf nitrogen in electron transport (FB) and in Rubisco (FR) were relatively constant within the canopy. Leaf aging led to increased MA, larger or constant mass-based N content, larger phosphorous (P) and structural carbon contents, but decreased movable cation contents. Age-dependent increases in MA and N per dry mass meant that Jmax and Vcmax per area were weakly related to leaf age, with a trend of decreasing values in older leaves. However, Jmax and Vcmax per unit dry mass decreased 4-fold across the range of leaf age, primarily owing to decreases in apparent N investments in photosynthetic machinery. This decrease in apparent N investments in photosynthetic machinery was possibly the result of a larger fraction of N bound to cell walls, or of an enhanced CO2 diffusion resistance from the outer surface of cell walls to the chloroplasts in older leaves with thicker and more lignified cell walls. The age-dependent variation in apparent fractional investments of N in photosynthetic machinery reduced the generality of leaf nitrogen v. photosynthesis relationships. Photosynthetic characteristics qualitatively fitted the same patterns with leaf age in all species, but at a common leaf age, area-based leaf photosynthetic potentials depended on species-specific values of MA. These data collectively demonstrate important canopy and age-dependent controls on leaf structure, chemistry and photosynthetic potentials that should be included in larger-scale photosynthesis simulations in Mediterranean climates.
Keywords: age effects, leaf structure, nitrogen v. photosynthesis relationship, nutrient content, sclerophyll species.
Acknowledgments
We thank Annette Suske for the help with chemical analyses. Financial support was provided by the Estonian Science Foundation (grant 5702), the Estonian Ministry of Education and Science (grant 0182468As03) and the Deutsche Bundesministerium für Bildung, Wissenschaft, Forschung und Technologie (Grant PT BEO 51 0339476B).
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