Structural controls on photosynthetic capacity through juvenile-to-adult transition and needle ageing in Mediterranean pines

1. Needle photosynthetic potentials strongly vary among primary (juvenile) and secondary (adult) needles (heteroblasty) in Pinus species, but there is limited understanding of the underlying structural, diffusional and chemical controls. 2. We studied differences in needle photosynthetic characteristics among current-year juvenile and adult needles and among different-aged adult needles in Mediterranean pines Pinus halepensis Mill., P.pinea L. and P.nigra J. F. Arnold subsp. salzmannii (Dunal) Franco, hypothesizing that needle anatomical modifications upon juvenile-to-adult transition lead to reduced photosynthetic capacity due to greater limitation of photosynthesis by mesophyll conductance and due to an increase in the share of support tissues at the expense of photosynthetic tissues. We also hypothesized that such alterations occur with needle ageing, but to a lower degree. 3. Photosynthetic capacity per dry mass was 2.4- to 2.7-fold higher in juvenile needles, and this was associated with 3.4- to 3.7-fold greater mesophyll diffusion conductance, 2- to 2.5-fold greater maximum carboxylase activity of Rubisco (V-cmax) and 2.2- to 3-fold greater capacity for photosynthetic electron transport (J(max)). The latter differences were driven by modifications in mesophyll volume fraction and changes in the share of nitrogen between structural and photosynthetic functions. Analogous changes in photosynthetic characteristics occurred with needle ageing, but their extent was less. 4. These results indicate that conifer foliage photosynthetic machinery undergoes a profound change from a fast return strategy in juveniles to slow return stress-resistant strategy in adults and that this strategy shift is driven by modifications in foliage biomass investments in support and photosynthetic functions as well as by varying mesophyll diffusional controls on photosynthesis. Changes in needle morphophysiotype during tree and needle ageing need consideration in predicting changes in tree photosynthetic potentials through tree ontogeny and during and among growing seasons.