Coexistent Heteroblastic Needles of Adult Pinus canariensis C.Sm. ex DC. in Buch Trees Differ Structurally and Physiologically

Great variation in shape and size between primary (juvenile) and secondary (adult) needles, so-called leaf-heteroblasty, occurs in several Pinus species. Most of them loss primary needles during the juvenile-to-adult transition of the tree. An exception to this is Pinus canariensis (a Canary Islands endemism) in which basal resprouting twigs of adult trees frequently wear both primary and secondary needles. Taking advantage of this extraordinary study-case-species, we conducted an exhaustive comparison of both needle types through quantitative analyses of needle anatomy, photochemical performance, gas exchange, and resistance to extreme dehydration and to extreme needle temperature. We hypothesized that primary needles would show lower investment to leaf structure but higher photosynthetical efficiency. Primary needles had less stomatal density and thicker and less wettable cuticles. In cross section, primary needles showed smaller structural fraction (e.g., percent of hypodermis, endodermis and vascular tissue) and higher fraction of photosynthetic parenchyma. Significant differences between primary and secondary needles were not found in net carbon assimilation not in their leaf mass area values. Interestingly, secondary needles showed higher electron transport rate, and they were additionally much more efficient in retaining water under severe and controlled desiccant conditions. When subjected to extreme temperatures (-10 degrees to +50 degrees C), primary needles recovered better their photochemical efficiency than secondary needles, after +46 degrees and +48 degrees C heat-shock treatments. Our results indicate that both needle types broaden the diversity of physiological responses against environmental constrains in basal twigs of adult P. canariensis trees. Considering that this is a fire-resistant and resprouting species, this advantage could be particularly useful after a drastic environmental change such a fire or a gap opening in the forest.