Plastic responses of Abies pinsapo xylogenesis to drought and competition.

Radial growth and xylogenesis were studied to investigate the influence of climate variability and intraspecific competition on secondary growth in Abies pinsapo Boiss., a relic Mediterranean fir. We monitored the responses to three thinning treatments (unthinned control -C-, 30% -T30- and 60% -T60- of basal area removed) to test the hypothesis that they may improve the adaptation capacity of tree growth to climatic stress. We also assessed whether xylogenesis was differentially affected by tree-to-tree competition. Secondary growth was assessed using manual band dendrometers from 2005 to 2007. In 2006, xylogenesis (phases of tracheid formation) was also investigated by taking microcores and performing histological analyses. Seasonal dynamics of radial increment were modeled using Gompertz functions and correlations with microclimate and radiation were performed. Histological analyses revealed it as fundamental to calibrate the dendrometer estimates of radial increment and to establish the actual onset and end dates of tracheid production. The lower radial-increment rates and number of produced tracheids were observed in the trees subjected to high competition in the unthinned plots. The growing season differed among the plots, and its duration ranged from an average of 78 days in unthinned plots to 115 days in thinned ones (T60). Variations in the beginning of the growing season (13 April to 22 May) and earlywood-latewood transition (early August) were mainly determined by the temperature pattern, while the onset and the end of the growing season were related to both annual precipitation and tree-to-tree competition. The tracheid-formation phases of radial enlargement and cell-wall thickening showed similar patterns in the trees from thinned and unthinned plots subjected to low and high competition, respectively, but the mean number of tracheids in each phase was always higher in the trees from the thinned plots. The reduction of competition through thinning induced a longer growing season and enhanced the radial growth in A. pinsapo.