Performance of two Picea abies (L.) Karst. stands at different stages of decline : VII. Nutrient relations and growth

A declining, closed-canopy Picea abies (L.) Karst. stand produced as much crown biomass as a healthy stand, although some trees were chlorotic due to magnesium deficiency. The production of wood per unit of leaf area in both stands was related to the foliar magnesium concentration. Although leaf area index and climate were similar at both sites, stemwood production was 35% lower in the declining than in the healthy stand. Nutritional disharmony, rather than a deficiency in a single element, was identified as the mechanism for reduced tree vigor. The role of nutrient stress in forest decline was detected by partitioning the season into three periods reflecting different phenological stages: a canopy growth period in spring, a stem growth period in summer, and a recharge period during the non-growing season. Needle growth was associated with nitrogen supply. Most of the magnesium supply required to meet the demand for foliage growth was retranslocated from mature needles. Magnesium retranslocation was related to concentration of nitrogen and magnesium in those needles before bud break. Retranslocation from mature needles during the phase of canopy production resulted in chlorosis in initially green needles if the magnesium concentration before bud break was low. Nitrogen concentration in 0-year-old needles generally remained constant with increasing supply, indicating that foliage growth was restricted by the supply of nitrogen. In contrast, magnesium concentration generally increased with supply, indicating that magnesium supply for needle growth was sufficient. Much of the magnesium required for wood production was taken up from the soil because stored magnesium was largely used for canopy growth. Uptake at the declining site was probably limited because of restricted root expansion and lower soil magnesium compared to the healthy site. For this reason only wood growth was reduced at the declining site. Because the recharge of magnesium during the non-growing period is dependent on uptake from the soil, it was more limited at the declining that at the healthy stand. However, as nitrogen uptake from the atmosphere may account for an appreciable proportion of the total uptake, and as its supply in the soil at both sites was similar, an unbalanced recharge of nitrogen and magnesium may have occurred at the declining site. If mature needles are unable to recharge with magnesium in proportion to the uptake of nitrogen, chlorosis is likely to occur during the next canopy growth period.