Population differentiation in climate sensitivity of resin duct formation during growth resumption in Pinus pinaster

To counteract the effects of herbivores and pathogens, conifers have developed a sophisticated resin-based defensive system. Since defences are costly, trees must continuously accommodate defensive investment throughout plastic responses to environmental stimuli. However, the extent of such responses can differ at the intra-specific level (i.e. genetic variation in plasticity). Here we examined whether and to what extent year-to-year climate fluctuations, an important source of environmental heterogeneity during the trees' life, drive plasticity in defensive allocation of a widespread pine species. Specifically, we quantified interannual variation in resin duct production along a 31-year-period in 174 Pinus pinaster trees of nine range-wide populations grown in two common gardens in Central Spain. We aimed to explore (i) patterns of interannual variation (i.e., temporal plasticity) in resin duct production among populations and sites, (ii) whether such patterns are linked to plastic responses to interannual variation in climate conditions (i.e., climatic plasticity), and (iii) whether plastic responses to climate differ among populations (i.e., genetic variation in plasticity) and sites. We found large interannual plasticity in resin duct production (22.8 % of total variance), with temporal patterns differing among sites and populations. Climate conditions during the early growth period significantly affected the annual differentiation of resin ducts. Particularly, April precipitation had a positive overall effect on resin duct production. Inversely, warmer conditions in April had a negative effect but only in certain populations, which demonstrates genetic variation in climate sensitivity of resin duct formation. Despite significant effects of certain climate variables on annual resin duct production, climate only accounted for a small proportion of the total interannual variation (up to 3.8 % of interannual variation explained by climate factors). This suggests that alternative factors such as trade-offs with growth and temporal variation in biotic and non-climatic abiotic conditions likely contribute to explain interannual fluctuations in defensive investment.