To determine effects of stem nitrogen concentration ([N]) on the seasonal course of respiration, rates of stem respiration of ten control and ten irrigated-fertilized (IL), 30-year-old Norway spruce trees (Picea abies (L.) Karst.), growing in northern Sweden, were measured on seven occasions from June 1993 to April 1994. To explore sources of seasonal variation and mechanisms of fertilization effects on respiration, we separated total respiration into growth and maintenance respiration for both xylem and phloem bark. Stem respiration increased in response to the IL treatment and was positively correlated with growth rate, volume of living cells and stem nitrogen content. However, no significant effect of IL treatment or [N] in the living cells was found for respiration per unit volume of live cells. Total stem respiration during the growing season (June to September) was estimated to be 16.7 and 29.7 mol CO(2) m(-2) for control and IL-treated trees, respectively. Respiration during the growing season accounted for approximately 64% of total annual respiration. Depending on the method, estimated growth respiration varied between 40 and 60% of total respiration during the growing season. Between 75 and 80% of the live cell volume in the stems was in the phloem, and phloem maintenance accounted for about 70% of maintenance respiration. Because most of the living cells were found in the phloem, and the living xylem cells were concentrated in the outer growth rings, we concluded that the best base for expressing rates of stem growth and maintenance respiration in young Norway spruce trees is stem surface area.

To study the effects of elevated CO(2) on gas exchange, nonstructural carbohydrate and nutrient concentrations in current-year foliage of 30-year-old Norway spruce (Picea abies (L.) Karst.) trees, branches were enclosed in ventilated, transparent plastic bags and flushed with ambient air (mean 370 &mgr;mol CO(2) mol(-1); control) or ambient air + 340 &mgr;mol CO(2) mol(-1) (elevated CO(2)) during two growing seasons. One branch bag was installed on each of 24 selected trees from control and fertilized plots. To reduce the effect of variation among trees, results from each treated branch were compared with those from a control branch on the same whorl of the same tree. Elevated CO(2) increased rates of light-saturated photosynthesis on average by 55% when measured at the treatment CO(2) concentration. The increase was larger in shoots with high needle nitrogen concentrations than in shoots with low needle nitrogen concentrations. However, shoots grown in elevated CO(2) showed a decrease in photosynthetic capacity compared with shoots grown in ambient CO(2). When measured at the internal CO(2) concentration of 200 &mgr;mol CO(2) mol(-1), photosynthetic rates of branches in the elevated CO(2) treatments were reduced by 8 to 32%. The elevated CO(2) treatment caused a 9 to 20% reduction in carboxylation efficiency and an 18% increase in respiration rates. In response to elevated CO(2), starch, fructose and glucose concentrations in the needles increased on average 33%, whereas concentrations of potassium, nitrogen, phosphorus, magnesium and boron decreased. Needle nitrogen concentrations explained 50-60% of the variation in photosynthesis and CO(2) acclimation was greater at low nitrogen concentrations than at high nitrogen concentrations. We conclude that the enhanced photosynthetic rates found in shoots exposed to elevated CO(2) increased carbohydrate concentrations, which may have a negative feedback on the photosynthetic apparatus and stimulate cyanide-resistant respiration. We also infer that the decrease in nutrient concentrations of needles exposed to elevated CO(2) was the result of retranslocation of nutrients to other parts of the branch or tree.

Few studies have examined variation in respiration rates within trees, and even fewer studies have focused on variation caused by within-stem temperature differences. In this study, stem temperatures at 40 positions in the stem of one 30-year-old Norway spruce (Picea abies (L.) Karst.) were measured during 40 days between July 1994 and June 1995. The temperature data were used to simulate variations in respiration rate within the stem. The simulations assumed that the temperature-respiration relationship was constant (Q10 = 2) for all days and all stem positions. Total respiration for the whole stem was calculated by interpolating the temperature between the thermocouples and integrating the respiration rates in three dimensions. Total respiration rate of the stem was then compared to respiration rate scaled up from horizontal planes at the thermocouple heights (40, 140, 240 and 340 cm) on a surface area and on a sapwood volume basis. Simulations were made for three distributions of living cells in the stems: one with a constant 5% fraction of living cells, disregarding depth into the stem; one with a living cell fraction decreasing linearly with depth into the stem; and one with an exponentially decreasing fraction of living cells. Mean temperature variation within the stem was 3.7 degrees C, and was more than 10 degrees C for 8% of the time. The maximum measured temperature difference was 21.5 degrees C. The corresponding mean variation in respiration was 35% and was more than 50% for 24% of the time. Scaling up respiration rates from different heights between 40 and 240 cm to the whole stem produced an error of 2 to 58% for the whole year. For a single sunny day, the error was between 2 and 72%. Thus, within-stem variations in temperature may significantly affect the accuracy of scaling respiration data obtained from small samples to whole trees. A careful choice of chamber position and basis for scaling is necessary to minimize errors from variation in temperature.

AbstractmRespiration of 1-year-old needles of 30-year-old Norway spruce trees [Picea abies (L.) Karst.] was studied in a nutrient optimisation experiment in northern Sweden. Respiration rates of detached needles, from ten control (C) and ten irrigated-fertilised (IL) trees, were measured on 16 occasions from June 1992 to June 1993. The aim of the study was to determine the influence of temperature on the seasonal course of needle maintenance respiration, and the effect of nitrogen concentration [N] and carbohydrate content on needle respiration in young Norway spruce trees subjected to long-term fertilisation. The IL treatment significantly affected needle size, in terms of dry mass and length, but not specific needle length (SNL). There was, however, a strong tree-specific effect on SNL (P510 ±9 , R 2 = 0.75). Needle starch content varied markedly with season (0±25% of total dry mass). This, unless accounted for, would cause erroneous estimates of nutrient concentrations, and of rates of needle respiration, within and between treatments. There was considerable seasonal variation in needle respiration, both in terms of maintenance respiration and temperature dependence (Q10). Q10 had its highest value (2.8) during winter and its lowest (2.0) in the middle of summer. In early autumn (August, September), respiration rate and needle [N] were significantly related (C: P = 0.001, IL: P50.0005). There was no significant difference in the slope between the two regression lines, but a difference in intercept. At the same needle [N], needles from IL-plots always had a lower respiration rate than needles from control plots. No obvious explanation for the observed difference in intercept was found, but some plausible assumptions are put forward and discussed.