Significance: Forest trees can live for hundreds to thousands of years, and they play a critical role in mitigating global warming by fixing approximately 15% of anthropogenic CO ₂ emissions annually by wood formation. However, the environmental factors triggering wood formation onset in springtime and the cellular mechanisms underlying this onset remain poorly understood, since wood forms beneath the bark and is difficult to monitor. We report that the onset of wood formation in Northern Hemisphere conifers is driven primarily by photoperiod and mean annual temperature. Understanding the unique relationships between exogenous factors and wood formation could aid in predicting how forest ecosystems respond and adapt to climate warming, while improving the assessment of long-term and high-resolution observations of global biogeochemical cycles.

Non-structural carbohydrates (NSCs) play a crucial role in xylem formation and represent, with water, the main constraint to plant growth. We assessed the relationships between xylogenesis and NSCs in order to (1) verify the variance explained by NSCs and (2) determine the influence of intrinsic (tissue supplying carbon) and extrinsic (water availability and temperature) factors. During two years, wood formation was monitored in saplings of black spruce submitted to a dry period of about one month in June and exposed to different temperature treatments in a greenhouse. In parallel, NSCs concentrations were determined by extracting the sugar compounds from two tissues (cambium and inner xylem), both potentially supplying carbon for wood formation. A mixed-effect model was used to assess and quantify the potential relationships. Total xylem cells, illustrating meristematic activity, were modeled as a function of water, sucrose and pinitol (conditional R2 of 0.79). Water availability was ranked as the most important factor explaining total xylem cell production, while the contribution of carbon was lower. Cambium stopped dividing under water deficit, probably to limit the number of cells remaining in differentiation without an adequate amount of water. By contrast, carbon factors were ranked as the most important in explaining the variation in the living cells (conditional R2 of 0.49) highlighting the functional needs during xylem development, followed by the tissue supplying the NSCs (cambium) and water availability. This study precisely demonstrates the role of carbon and water in structural growth expressed as meristematic activity and tissue formation.