Energetics of maize C4 physiology under light limiting conditions
C4 plants have a biochemical carbon concentrating mechanism (CCM) that increases CO2 concentration around Rubisco in the bundle sheath (BS). Maize CCM has two CO2 delivery pathways to the Bundle Sheath (BS) (respectively via malate, MAL or aspartate, ASP); rates of PGA reduction, carbohydrate synthesis and PEP regeneration vary between BS and Mesophyll (M) cells. For these anatomical and biochemical complexities, C4 plants are highly sensitive to light conditions. Under limiting light, the activity of the CCM generally decreases, causing an increase in leakiness, (Φ), the ratio of CO2 retrodiffusing from the BS relative to C4 carboxylation processes. This increase in Φ had been theoretically associated with a decrease in biochemical operating efficiency (expressed as ATP cost of gross assimilation, ATP / GA) under low light and, because a proportion of canopy photosynthesis is carried out by shaded leaves, to potential productivity losses at field scale. In C4 leaves, because of the concentric anatomy, light reaches M cells before the deeper BS (Evans et al., 2007), and could alter the energetic partitioning balance between BS and M and potentially cause efficiency losses. In this experimental programme I investigated strategies deployed by C4 plants to adjust operating efficiency under different illumination conditions. Firstly, maize plants were grown under high and low light regimes (respectively HL, 600 vs LL, 100 μE m-2 s-1). Short term acclimation of Φ was compared from isotopic discrimination (Δ), gas exchange and photochemistry using an improved modelling approach which does not suffer from elements of circularity. Long term acclimation to low light intensities brought about physiological changes which could potentially increase the operating efficiency under limiting ATP supplies. Secondly, profiles of light penetration across a leaf were used to derive the potential ATP supply for M and BS cells induced by changing light quality. Empirical measurements of net CO2 uptake, ATP production rate and carbon isotope discrimination were made on plants under a low light intensity. The overall conversion efficiency was not affected by light quality. A comprehensive metabolic model highlighted the importance of both CO2 delivery pathways in maize. Further, metabolic plasticity allowed the balancing of ATP and NADPH requirements between BS and M. Finally, I tested the hypothesis that plants can modify their physiology so as to reach a status of higher operating efficiency when exposed to high light and then to low light, so as to mimic the transition which leaves undergo when shaded by newly emerging leaves in a crop canopy. Plants were grown under high light and low light for three weeks, then, HL plants were transferred to low light for a further three weeks. Re-acclimation was very effective in reducing ATP cost of net assimilation under low light intensities. In addition, the hyperbolic leakiness increase observed under low light intensities was not associated to operating efficiency loss. Overall, in the three experimental Chapters I showed compelling theoretical and empirical evidence proving the hypothesis that C4 plants deal with low light conditions and with different light qualities without losing operating efficiency.
photosynthesis, carbon reduction, energetics, shade, C4, light quality, biochemistry, light penetration, model, carbon isotope discrimination, leakiness, bundle sheath
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