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JXB Advance Access originally published online on December 15, 2005
Journal of Experimental Botany 2006 57(2):343-354; doi:10.1093/jxb/erj014
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© The Author [2005]. Published by Oxford University Press [on behalf of the Society for Experimental Biology]. All rights reserved. For Permissions, please e-mail: journals.permissions@oxfordjournals.org

RESEARCH PAPER

Irradiance and phenotype: comparative eco-development of sun and shade leaves in relation to photosynthetic CO2 diffusion

Ichiro Terashima1,*, Yuko T. Hanba2, Youshi Tazoe1, Poonam Vyas1 {dagger} and Satoshi Yano1 {ddagger}

1Department of Biology, Graduate School of Science, Osaka University, 1-1 Machikaneyama-cho, Toyonaka, 560-0043, Japan
2Center for Bioresource Field Science, Kyoto Institute of Technology, 1 Saga-ippongi-cho, Ukyo-ku, Kyoto, 616-8354, Japan

* To whom correspondence should be addressed. E-mail: itera{at}bio.sci.osaka-u.ac.jp

The subject of this paper, sun leaves are thicker and show higher photosynthetic rates than the shade leaves, is approached in two ways. The first seeks to answer the question: why are sun leaves thicker than shade leaves? To do this, CO2 diffusion within a leaf is examined first. Because affinity of Rubisco for CO2 is low, the carboxylation of ribulose 1,5-bisphosphate is competitively inhibited by O2, and the oxygenation of ribulose 1,5-bisphosphate leads to energy-consuming photorespiration, it is essential for C3 plants to maintain the CO2 concentration in the chloroplast as high as possible. Since the internal conductance for CO2 diffusion from the intercellular space to the chloroplast stroma is finite and relatively small, C3 leaves should have sufficient mesophyll surfaces occupied by chloroplasts to secure the area for CO2 dissolution and transport. This explains why sun leaves are thicker. The second approach is mechanistic or ‘how-oriented’. Mechanisms are discussed as to how sun leaves become thicker than shade leaves, in particular, the long-distance signal transduction from mature leaves to leaf primordia inducing the periclinal division of the palisade tissue cells. To increase the mesophyll surface area, the leaf can either be thicker or have smaller cells. Issues of cell size are discussed to understand plasticity in leaf thickness.

Key words: Aquaporin, cell wall, chloroplasts, conductance, diffusion, intercellular spaces, mechanical strength, photosynthesis, resistance to CO2 diffusion, stomata


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