Drought will increasingly threaten forest ecosystems worldwide. Understanding how competition influences tree growth response to drought is essential for forest management aiming at climate change adaptation. However, published results from individual case studies are heterogeneous and sometimes contradictory. We reviewed 166 cases from the peer-reviewed literature to assess the influence of stand-level competition on tree growth response to drought. We monitored five indicators of tree growth response: mean sensitivity (inter-annual tree ring width variability); association between inter-annual growth variability and water availability; resistance; recovery; and resilience to drought. Vote counting did not indicate a consistent effect of competition on mean sensitivity. Conversely, higher competition for resources strengthened the association between water availability and inter-annual growth rates. Meta-analysis showed that higher competition reduced resistance (p < 0.001) and improved recovery (p < 0.05), but did not consistently affect resilience. Species, site and stand characteristics, and drought intensity were insignificant or poor predictors for the large variability among the investigated cases. Our review and meta-analysis show that competition does not affect the response of tree growth to drought in a unidirectional and universal way. Although density reduction (thinning) can alleviate growth declines during drought, the effects on growth after stress are uncertain. The large variability among investigated cases suggests that local-scale processes play a crucial role in determining such responses and should be explicitly evaluated and integrated into specific strategies for adaptation of forests to climate change.

Background and Aims During the growing season, the cambium of conifer trees produces successive rows of xylem cells, the tracheids, that sequentially pass through the phases of enlargement and secondary wall thickening before dying and becoming functional. Climate variability can strongly influence the kinetics of morphogenetic processes, eventually affecting tracheid shape and size. This study investigates xylem anatomical structure in the stem of Picea abies to retrospectively infer how, in the long term, climate affects the processes of cell enlargement and wall thickening. Methods Tracheid anatomical traits related to the phases of enlargement (diameter) and wall thickening (wall thickness) were innovatively inspected at the intra-ring level on 87-year-long tree-ring series in Picea abies trees along a 900 m elevation gradient in the Italian Alps. Anatomical traits in ten successive tree-ring sectors were related to daily temperature and precipitation data using running correlations. Key Results Close to the altitudinal tree limit, low early-summer temperature negatively affected cell enlargement. At lower elevation, water availability in early summer was positively related to cell diameter. The timing of these relationships shifted forward by about 20 (high elevation) to 40 (low elevation) d from the first to the last tracheids in the ring. Cell wall thickening was affected by climate in a different period in the season. In particular, wall thickness of late-formed tracheids was strongly positively related to August–September temperature at high elevation. Conclusions Morphogenesis of tracheids sequentially formed in the growing season is influenced by climate conditions in successive periods. The distinct climate impacts on cell enlargement and wall thickening indicate that different morphogenetic mechanisms are responsible for different tracheid traits. Our approach of long-term and high-resolution analysis of xylem anatomy can support and extend short-term xylogenesis observations, and increase our understanding of climate control of tree growth and functioning under different environmental conditions.

Understanding how climate affects xylem formation is critical for predicting the impact of future conditions on tree growth and functioning in the Mediterranean region, which is expected to face warmer and drier conditions. However, mechanisms of growth response to climate at different temporal scales are still largely unknown, being complicated by separation between spring and autumn xylogenesis (bimodal temporal pattern) in most species such as Mediterranean pines. We investigated wood anatomical characteristics and carbon stable isotope composition in Mediterranean Pinus pinea L. along tree-ring series at intra-ring resolution to assess xylem formation processes and responses to intra-annual climate variability. Xylem anatomy was strongly related to environmental conditions occurring a few months before and during the growing season, but was not affected by summer drought. In particular, the lumen diameter of the first earlywood tracheids was related to winter precipitation, whereas the size of tracheids produced later was influenced by mid-spring precipitation. Diameter of latewood tracheids was associated with precipitation in mid-autumn. In contrast, tree-ring carbon isotope composition was mostly related to climate of the previous seasons. Earlywood was likely formed using both recently and formerly assimilated carbon, while latewood relied mostly on carbon accumulated many months prior to its formation. Our integrated approach provided new evidence on the short-term and carry-over effects of climate on the bimodal temporal xylem formation in P. pinea. Investigations on different variables and time scales are necessary to disentangle the complex climate influence on tree growth processes under Mediterranean conditions.

Climate change can induce substantial modifications in xylem structure and water transport capacity of trees exposed to environmental constraints. To elucidate mechanisms of xylem plasticity in response to climate, we retrospectively analysed different cell anatomical parameters over tree-ring series in Norway spruce (Picea abies L. Karst.). We sampled 24 trees along an altitudinal gradient (1200, 1600 and 2100 m above sea level, a.s.l.) and processed 2335 ± 1809 cells per ring. Time series for median cell lumen area (MCA), cell number (CN), tree-ring width (RW) and tree-ring-specific hydraulic conductivity (Kr) were crossed with daily temperature and precipitation records (1926–2011) to identify climate influence on xylem anatomical traits. Higher Kr at the low elevation site was due to higher MCA and CN. These variables were related to different aspects of intra-seasonal climatic variability under different environmental conditions, with MCA being more sensitive to summer precipitation. Winter precipitation (snow) benefited most parameters in all the sites. Descending the gradient, sensitivity of xylem features to summer climate shifted mostly from temperature to precipitation. In the context of climate change, our results indicate that higher summer temperatures at high elevations will benefit cell production and xylem hydraulic efficiency, whereas reduced water availability at lower elevations could negatively affect tracheids enlargement and thus stem capacity to transport water.