Why does photosynthesis decrease with needle age in Pinus pinaster?

By C. Warren. Trees, 2006

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Mesophyll conductance to CO2: current knowledge and future prospects. (2008, methodology): “Warren (2006a) reported that SDs could be as high as 19–38% of the mean for the chlorophyll fluorescence methods, and 6–34% for the isotopic method.”
The role of crown architecture, leaf phenology and photosynthetic activity in promoting complementary use of light among coexisting species in temperate forests (2010, background): “Photosynthetic activity also varies with leaf age (e.g., Reich et al. 2009; Warren 2006), such that leaves of evergreen understory trees may mature later in the season so that their photosynthetic capacity is high when leaves of the deciduous canopy trees begin to shed in autumn.”
Ecology of Leaf Longevity (2011, background): “Even if the microenvironment around a leaf is stable over its lifetime, cumulative damage and reduced internal conductance of CO 2 (Hensel et al. 1993; Guarente et al. 1998; Nooden 2004; Warren 2006) can lead to gradually lower photosynthetic capacity in older leaves.”