However, information about these metabolic and physiological processes is very
limited.
1.8.1 Carbon Metabolism (Fig. 1.20)
• 1st step: Solar radiation energy (electron) is transported through photosystem II
(PSII) and photosystem I (PSI) in chloroplasts, and high-energy compounds, ATP
and NADPH, are produced.
• 2nd step: Solar radiation energy stimulates the K
+ pump to open stomata with
osmotic pressure, and CO 2 and H 2 O diffuse into and out of the cell, respectively. As stomata are porous structures, when H2O evaporates as H 2 O gas due
to solar radiation energy, the vaporization heat (latent heat of vaporization) is
removed, and the leaf is cooled. As CO 2 and H 2 O diffusion are physical
processes powered by solar radiation, CO 2 and H 2 O diffusion are coupled.
This relationship is called water-use efficiency (WUE), defined as the amount
of carbon assimilated as biomass or grain produced per unit of water used by the
plant (WUE ¼ P/E ¼ dry matter (DM) production g/water g), where P and E are
the photosynthetic rate and the evaporation rate, respectively. The WUE
(g DM/liter water) (https://photosyn.jp/pwiki/index.php?%E6%B0%B4%E5%
88%A9%E7%94%A8%E5%8A%B9%E7%8E%87) is 2.5–5 in trees and
1.4–2.5 in grasses, which are equivalent in g water/g DM to 200–400 in trees
and 400–700 in grasses. These values indicate that photosynthetic processes
require large amounts of water and that the WUE of trees is higher than that of
grasses.
• 3rd step: CO 2 is reduced to triose-P through the Calvin cycle using energy from
ATP and NADPH
• 4th step: Triose-P is metabolized into sucrose (carbohydrate-oriented metabolism) and PEP-OAA (organic acid- and amino acid-oriented metabolism), the
regulation of which is mainly by SPS and PECP. PECP is activated by light (solar
radiation), NO 3
À , and Pi (indirectly).
• 5th step: PEP is retranslocated into chloroplasts regulated by PPT (PPT: PEP/Pi
translocator), and PEP is metabolized into precursors of high molecular weight
compounds such as pigments, lignin, aromatic amino acids, and fatty acids. As
trees produce a large amount of lignin, this metabolic pathway is very important.
The production of precursors to high molecular weight compounds is assumed to
require large amounts of energy; however, solar energy is abundant in tropical
areas, and chemical construction energy is provided inside the chloroplasts,
contributing to the consumption of excess energy.
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M. Osaki et al.
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