Gas exchange and leaf aging in an evergreen oak: causes and consequences for leaf carbon balance and canopy respiration.

Leaves of Mediterranean evergreens experience large variations in gas exchange rates over their life span due to aging and seasonally changing environmental conditions. Accounting for the changing respiratory physiology of leaves over time will help improve estimations of leaf and whole-plant carbon balances. Here we examined seasonal variations in light-saturated net CO(2) assimilation (A(max)), dark respiration (R(d)) and the proportional change in R(d) per 10 °C change in temperature (Q(10) of R(d)) in previous-year (PY) and current-year (CY) leaves of the broadleaved evergreen tree Quercus ilex L. A(max) and R(d) were lower in PY than in CY leaves. Differences in nitrogen between cohorts only partly explained such differences, and rates of A(max) and R(d) expressed per unit of leaf nitrogen were still significantly different between cohorts. The decline in A(max) in PY leaves did not result in the depletion of total non-structural carbohydrates, whose concentration was in fact higher in PY than CY leaves. Leaf-level carbon balance modeled from gas exchange data was positive at all ages. Q(10) of R(d) did not differ significantly between leaf cohorts; however, failure to account for distinct R(d) between cohorts misestimated canopy leaf respiration by 13% across dates when scaling up leaf measurements to the canopy. In conclusion, the decline in A(max) in old leaves that are close to or exceed their mean life span does not limit the availability of carbohydrates, which are probably needed to sustain new growth, as well as R(d) and nutrient resorption during senescence. Accounting for leaf age as a source of variation of R(d) improves the estimation of foliar respiratory carbon release at the stand scale.