Carbon Isotope Discrimination and Photosynthetic Gas Exchange in Coffee Hedgerows During Canopy Development

Abstract

In evergreen species, leaf carbon isotope discrimination (Δ) integrates phenological rhythms in gas exchange as well as seasonal changes in environmental conditions. However, few reports on long term variations in Δ of woody plants are available. We measured Δ, gas exchange, nitrogen content, and photosynthetic nitrogen-use efficiency (PNUE) in coffee hedgerows at different stages of canopy development encompassing a range of leaf area index (LAI) from 0.7 to 7.5. Assimilation was highest in sun leaves, but stomatal conductance was highest in shaded leaves. This resulted in a high correlation between assimilation and stomatal conductance in sun, but not in shaded leaves. Δ was about 20 lower in sun than in shaded leaves, and varied by 2.30 among leaves at different positions along two-year- old branches. These differences in Δ were the result of changes in carbon isotope composition that occurred in mature, fully expanded leaves as they became shaded during subsequent canopy growth. Results from a mass balance model based on leaf gas exchange characteristics and measured foliar Δ values suggested that about 50% of the carbon originally fixed during leaf development in the sun may have subsequently been turned over in the shade. Δ of sun leaves from the upper canopy decreased by about 20 with increasing LAI, indicating that intrinsic water-use efficiency (WUE) of this canopy layer increased during canopy development. In contrast, instantaneous WUE, estimated as assimilation divided by canopy transpiration obtained from sap flow measurements, seemed to decrease with increasing LAI. PNUE of upper canopy sun leaves decreased with increasing LAI, suggesting a physiological compromise between WUE and PNUE mediated by stomatal conductance, which also decreased with increasing LAI. A strong negative correlation obtained between leaf Δ and N content was consistent with a trade-off between intrinsic water- and N-use efficiency.