The intra-annual rhythm of Pinus sylvestris growth-climate responses under a warming climate at its southern distribution limits

Warming climate has posed a pressing threat on boreal forests and an improved understanding of the intra-annual pattern of climatic influences on the tree growth can help interpret the response of boreal forest to climate change. Here, we systematically examined the growth-climate relationship, cambium phenology, and xylem cell dynamics of Pinus sylvestris (PS) to disentangle its intra-annual growth rhythm to external drivers and internal physiological process at its southern distribution limits across Eurasia. We showed the intra-annual rhythm of PS to temperature and precipitation is anti-phase and synchronous at its southern limits. Temperature had both promoting and inhibiting effects on PS growth over growing season where the growth-temperature response gradually changed from positive in winter-early spring to negative in late spring-summer, and then returned to positive in autumn. Precipitation enhanced PS growth in late spring-summer. Temperature dominated the intra-annual growth-climate response rhythm, and the response shifted to negative when temperature reached 13.72 °C in spring-summer and shifted to positive when temperature reached 17.41 °C in summer-autumn in Shenyang, respectively. A high water demand for the rapid earlywood cell formation in spring-summer and heat requirement for cambial cell division in summer-autumn caused the shifts in growth-temperature response, respectively. Climatic warming advances and prolongs the time of summer water availability that limits PS growth at its southern distribution limits and thus warming climate may pose a greater threat to the southern population of PS.