In the Nordic countries, growth of Norway spruce (Picea abies (L.) Karst.) is generally limited by low availability of nutrients, especially nitrogen. Optimizing forest management requires better insight on how growth responds to the environmental conditions and their manipulation. The aim of this study was to analyse the effects of nutrient optimization on timing and the rate of tracheid formation of Norway spruce and to follow the differentiation of newly formed tracheids. The study was performed during two growing seasons in a long-term nutrient optimization experiment in northern Sweden, where all essential macro- and micronutrients were supplied in irrigation water every second day from mid-June to mid-August. The control plots were without additional nutrients and water. Tracheid formation in the stem was monitored throughout the growing season by weekly sampling of microcores at breast height. The onset of xylogenesis occurred in early June, but in early summer there were no significant between-treatment differences in the onset and relative rate of tracheid formation. In both treatments, the onset of secondary cell wall formation occurred in mid-June. The maximum rate of tracheid formation occurred close to the summer solstice and 50% of the tracheids had been accumulated in early July. Optimized nutrition resulted in the formation of ∼50% more tracheids and delayed the cessation of tracheid formation, which extended the tracheid formation period by 20-50%, compared with control trees. The increased growth was mainly an effect of enhanced tracheid formation rate during the mid- and later-part of the growing season. In the second year, the increased growth rate also resulted in 11% wider tracheids. We conclude that the onset and rate of tracheid formation and differentiation during summer is primarily controlled by photoperiod, temperature and availability of nutrients, rather than supply of carbohydrates.
Keywords: Picea abies; boreal forest; fertilization; microcores; phenology; temperature sum; tracheids; xylogenesis.