Abstract
Bifacial leaves contain an array of chloroplasts which are aligned along cell walls adjacent to intercellular airspaces. Cells beneath the adaxial (upper) surface are generally cylindrical in shape, forming palisade tissue and contain chloroplasts that have characteristics associated with acclimation to high irradiance. Conversely, cells near the abaxial (lower) surface are irregular in shape, forming spongy tissue and contain chloroplasts that have characteristics associated with acclimation to low irradiance. This results in a gradient of declining photosynthetic capacity per unit chlorophyll with increasing depth into the leaf. Space irradiance declines dramatically through the leaf. The profile of light absorption through the leaf has yet to be measured, but should approximate the product of space irradiance and pigment profiles. Since pigment content is highest in the middle of the leaf, light absorption peaks about one quarter of the way through the leaf. The response of leaf photosynthetic rate to irradiance reflects the interaction between the profiles of light absorption and chloroplast characteristics. Quantum yield (mol CO2 mol-1 quanta absorbed) varies for different layers in the leaf because the profile of Rubisco content per unit chlorophyll is not as steep as the light absorption profile. This skews the profile of carbon fixation away from the light absorption profile, favouring greater fixation in deeper layers. Nishio, Sun and Vogelmann (1993, Plant Cell 5, 953-961) measured profiles of 14C fixation across spinach leaves and concluded that carbon fixation was disconnected from the light gradient. Reanalysis of their data shows that carbon fixation is remarkably consistent with light absorption obeying the Beer- Lambert law when allowance is made for chlorophyll and Rubisco profiles through the leaf. This interpretation is supported by chlorophyll fluorescence properties of adaxial and abaxial surfaces. An apparent extinction coefficient of about 1500 m2 (mol Chl)-1 could be derived from leaf data which compares with 2230 m2 (mol Chl)-1 for pigment-protein complexes in solution. A possible test for this analysis would be to measure the 14C fixation profile of leaves illuminated on their abaxial surface.
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