Soil temperature,gas exchange and nitrogen status of 5-year-old Norway spruce seedlings

Five-year-old Norway spruce (Picea abies (L.) Karst.) seedlings were subjected to three simulated growing seasons in controlled environment chambers. Plants were acclimated to a soil temperature of 16 degrees C during the first and third growing seasons, but were allocated at random to soil temperature treatments of 9, 13, 18 and 21 degrees C during the second growing season. Low soil temperature during the second growing season depressed stomatal conductance and photosynthetic rate (A) per unit of projected leaf area, although intercellular CO2 concentrations did not differ significantly between treatments. At all soil temperatures, total chlorophyll concentration first decreased and then increased, although the rate of increase and the final concentration increased with soil temperature, which may explain the effect of soil temperature on A. Neither chlorophyll a/b ratio nor leaf nitrogen concentration was significantly affected by soil temperature. Treatment differences disappeared during the third simulated growing season when plants were again acclimated to a soil temperature of 16 degrees C.     

Short-day photoperiods affect expression of genes related to dormancy and freezing tolerance in Norway spruce seedlings

Key Message

Gene expression analysis showed that prolonged short day (SD) treatment deepened dormancy and stimulated development of freezing tolerance of Picea abies seedlings. Prolonged SD treatment also caused later appearance of visible buds in autumn, reduced risks for reflushing, and promoted earlier spring bud break.

Context

Short day (SD) treatment of seedlings is a common practice in boreal forest tree nurseries to regulate shoot growth and prepare the seedlings for autumn planting or frozen storage.

Aims

The aim of this study was to examine responses of Norway spruce (Picea abies (L.) Karst.) to a range of SD treatments of different length and evaluate gene expression related to dormancy induction and development of freezing tolerance.

Methods

The seedlings were SD treated for 11 h a day during 7, 14, 21, or 28 days. Molecular tests were performed, and the expression profiles of dormancy and freezing tolerance-related genes were analyzed as well as determination of shoot growth, bud set, bud size, reflushing, dry matter content, and timing of spring bud break.

Results

The 7-day SD treatment was as effective as longer SD treatments in terminating apical shoot growth. However, short (7 days) SD treatment resulted in later activation of dormancy-related genes and of genes related to freezing tolerance compared to the longer treatments which had an impact on seedling phenology.

Conclusion

Gene expression analysis indicated an effective stimulus of dormancy-related genes when the SD treatment is prolonged for at least 1–2 weeks after shoot elongation has terminated and that seedlings thereafter are exposed to ambient outdoor climate conditions.

     

Influence of nitrogen and phosphorous availability and ozone stress on Norway spruce seedlings

Four-year-old Norway spruce (Picea abies L. (Karst.)) seedlings were exposed to ambient and elevated (1.5 x ambient in 1997 and 1.6 x ambient in 1998) ozone concentrations [O3] and three nitrogen (N) and two phosphorus (P) availabilities: "optimal" values (control); 70% of the control N and P values (LN and LP); and 150% of the control N value (HN). Treatments were applied in an open-field ozone fumigation facility during the 1997 and 1998 growing seasons. Effects on growth, mineral and pigment concentrations, stomatal conductance and ultrastructure of needles were studied. The HN treatment increased growth significantly, whereas elevated [O3] had a slight or variable impact on growth and biomass allocation in all N treatments. Although there were no significant effects of the LP treatment on plant growth during the second year, there was a reduction in 1-year-old shoot dry mass in the elevated O3 + LP treatment at the end of the experiment. There were no significant treatment effects on mineral concentrations of current-year and 1-year-old needles at the final harvest. In response to the HN treatment, chlorophyll a and b and carotenoid concentrations increased significantly in current-year needles. Chlorophyll a/b ratio decreased in response to elevated [O3] alone, but increased in seedlings in the O(3) + LP treatment. Stomatal conductance of current-year needles decreased with increasing N availability, but increased in response to elevated [O3]. However, the O3-induced increase in stomatal conductance was less in the LN and LP treatments than in the control treatment. In chloroplasts of current-year needles, increased N availability decreased mean starch grain area, but increased the number of plastoglobuli. We conclude that Norway spruce seedlings are relatively tolerant to slightly elevated [O3], and that nitrogen and phosphorus imbalances do not greatly affect the influence of O3 on this species when the exposure lasts for two growing seasons or less.     

Impact of elevated carbon dioxide concentration and temperature on bud burst and shoot growth of boreal Norway spruce

Effects of elevated temperature and atmospheric CO2 concentration ([CO2]) on spring phenology of mature field-grown Norway spruce (Picea abies (L.) Karst.) trees were followed for three years. Twelve whole-tree chambers (WTC) were installed around individual trees and used to expose the trees to a predicted future climate. The predicted climate scenario for the site, in the year 2100, was 700 micromol mol-1 [CO2], and an air temperature 3 degrees C higher in summer and 5 degrees C higher in winter, compared with current conditions. Four WTC treatments were imposed using combinations of ambient and elevated [CO2] and temperature. Control trees outside the WTCs were also studied. Bud development and shoot extension were monitored from early spring until the termination of elongation growth. Elevated air temperature hastened both bud development and the initiation and termination of shoot growth by two to three weeks in each study year. Elevated [CO2] had no significant effect on bud development patterns or the length of the shoot growth period. There was a good correlation between temperature sum (day degrees>or=0 degrees C) and shoot elongation, but a precise timing of bud burst could not be derived by using an accumulation of temperature sums.     

Ectomycorrhizas and cadmium toxicity in Norway spruce seedlings

We studied the effects of ectomycorrhizal colonization by Laccaria bicolor (Maire) Orton S238 and Paxillus involutus (Batsch) Fr. 533 on cadmium (Cd) toxicity in Norway spruce seedlings (Picea abies (L.) Karst.). Both mycorrhizal and nonmycorrhizal seedlings were exposed to 0 (control), 0.5 or 5 &mgr;M CdSO(4) for 9 weeks in a sand culture system with frequent addition of nutrient solutions. In pure culture, P. involutus and L. bicolor showed similar Cd tolerance. However, in symbiosis, the Cd treatments decreased colonization by L. bicolor but not by P. involutus. Paxillus involutus ameliorated the negative effects of 0.5 &mgr;M Cd on shoot and root growth and chlorophyll content of old needles, whereas L. bicolor did not. Mycorrhizal colonization did not affect Cd concentrations of old needles and roots of seedlings. Despite differences between the ectomycorrhizal fungi in colonization and ability to alleviate Cd toxicity of seedlings, both species reduced Cd concentrations of young needles to a similar degree compared with nonmycorrhizal seedlings. However, in the 0.5 &mgr;M Cd treatment, the Cd content of needles of seedlings colonized by P. involutus was increased, whereas the Cd content of needles of seedlings colonized by L. bicolor was similar to that of needles of nonmycorrhizal seedings. When the amount of Cd translocated to needles was expressed on a root length basis to account for differences in the size of the root systems, the amount of Cd translocated to the needles was similar in seedlings mycorrhizal with P. involutus and in nonmycorrhizal seedlings. All mycorrhizal seedlings were similarly affected by 5 &mgr;M Cd, indicating that the amelioration efficiency of ectomycorrhizal fungi is dependent on the metal concentration to which the roots are exposed. Concentrations of P, K, Ca, Mg and Mn were decreased by 5 &mgr;M Cd to a similar extent in both nonmycorrhizal and mycorrhizal seedlings. In contrast to L. bicolor, P. involutus increased P uptake and altered patterns of root branching. We conclude that mycorrhizas alleviate Cd-induced reductions in growth of Picea abies seedlings. Although the two mycorrhizal fungi examined differed in their ability to alleviate Cd toxicity, these differences were not related to differences in Cd uptake or translocation to the shoot of the mycorrhizal seedlings. We suggest that amelioration of Cd toxicity by P. involutus may be a result of improved P nutrition or changes in root morphology, or both.     

Accentuation of gas exchange gradients in flushes of ponderosa pine exposed to ozone

Two-year-old ponderosa pine seedlings (Pinus ponderosa Laws.) were exposed to episodic O(3) concentrations in open-top chambers for two consecutive growing seasons (June through September of 1990 and 1991). Near the end of the second season of O(3) exposure, gas exchange was measured on needles of surviving flushes at saturating CO(2) and photosynthetic photon flux density (PPFD). Both photosynthetic capacity (A(sat)) and stomatal conductance to water vapor (g(wv)) declined linearly with needle age but differences within a flush were also found. Gas exchange rates of needles from the base of the current-year flush were significantly lower than those of needles from the top of the flush, even though age differences between needles were negligible. Although most measurements were conducted at saturating CO(2), similar patterns of gas exchange were also found at 350 micro mol mol(-1) CO(2), indicating that photosynthesis of needles at the bottom of the flush was more strongly affected by O(3) than that of needles at the top of the flush, even though the potential for O(3) uptake was probably less in needles at the bottom of the flush because of reduced stomatal conductance. Carboxylation efficiency (deltaA/deltaC(i)), the linear slope of the A/C(i) response, was highly correlated with A(sat), varying with needle age, needle position in the flush and O(3) exposure, but the magnitude of the reductions was greater than for A(sat). We conclude that susceptibility to O(3) damage among needles of an individual seedling varies not only with needle age but also with needle position, and that reductions in photosynthetic capacity may not be directly attributable to increased uptake of the pollutant. The data also indicate the need to consider within-flush variation when estimating whole-plant carbon gain and O(3) uptake.     

Effects of soil temperature and elevated atmospheric CO2 concentration on gas exchange, in vivo carboxylation and chlorophyll fluorescence in jack pine and white birch seedlings

One-year-old jack pine (Pinus banksiana Lamb.) and current-year white birch (Betula papyrifera Marsh.) seedlings were grown in ambient (360 ppm) or twice ambient (720 ppm) atmospheric CO2 concentration ([CO2]) and at three soil temperatures (Tsoil = 7, 17 and 27 degrees C initially, increased to 10, 20 and 30 degrees C two months later, respectively) in a greenhouse for 4 months. In situ foliar gas exchange, in vivo carboxylation characteristics and chlorophyll fluorescence were measured after 2.5 and 4 months of treatment. Low Tsoil suppressed net photosynthetic rate (Pn), stomatal conductance (g(s)) and transpiration rate (E) in jack pine in both CO2 treatments and g(s) and E in white birch in ambient [CO2], but enhanced instantaneous water-use efficiency (IWUE) in both species after 2.5 months of treatment. Treatment effects on g(s) and E remained significant throughout the 4-month study. Low Tsoil reduced maximal carboxylation rate (Vcmax) and PAR-saturated electron transport rate (Jmax) in jack pine in elevated [CO2] after 2.5 months of treatment, but not after 4 months of treatment. Low Tsoil increased actual photochemical efficiency of photosystem II (PSII) in the light (DeltaF/Fm') in jack pine, but decreased DeltaF/Fm' in white birch after 4 months of treatment. In response to low Tsoil, photosynthetic linear electron transport to carboxylation (Jc) decreased in jack pine after 2.5 months and in white birch after 4 months of treatment. Low Tsoil increased the ratio of the photosynthetic linear electron transport to oxygenation (Jo) to the total photosynthetic linear electron transport rate through PSII (Jo/J(T)) in both species after 2.5 months of treatment, but the effects became statistically insignificant in white birch after 4 months of treatment. High Tsoil decreased foliar N concentration in white birch. Elevated [CO2] increased Pn, IWUE and Jc but decreased Jo/J(T) in both species at both measurement times except Jc in white birch after 2.5 months of treatment. Elevated [CO2] also decreased g(s) and E in white birch at high Tsoil, Vcmax in both species and triose phosphate utilization in white birch at low Tsoil after 4 months of treatment, and DeltaF/Fm' in white birch after 2.5 months of treatment. Elevated [CO2] also increased foliar N concentration in both species. Low Tsoil caused no permanent damage to PSII in either species, but jack pine responded and acclimated to low Tsoil more quickly than white birch. Photosynthetic down-regulation and a decrease in photosynthetic electron transport to photorespiration occurred in both species in response to elevated [CO2].     

The effects of soil and air temperature on CO2 exchange and net biomass accumulation in Norway spruce, Scots pine and silver birch seedlings

Soil temperature is proposed to affect the photosynthetic rate and carbon allocation in boreal trees through sink limitation. The aim of this study was to investigate the effect of temperature on CO(2) exchange, biomass partitioning and ectomycorrhizal (ECM) fungi of boreal tree species. We measured carbon allocation, above- and below-ground CO(2) exchange and the species composition of associated ECM fungi in the rhizosphere of Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies K.) and silver birch (Betula pendula Roth) seedlings grown in soil maintained at 7-12, 12-15 and 16-22 °C. We found increased root biomass and photosynthetic rate at higher soil temperatures, but simultaneously with photosynthesis rate, higher temperature generally increased soil respiration as well as shoot, and root and rhizosphere respiration. The net CO(2) exchange and seedling biomass did not increase significantly with increasing temperature due to a concomitant increase in carbon assimilation and respiration rates. The 2-month-long growth period in different soil temperatures did not alter the ECM fungi species composition and the below-ground carbon sink strength did not seem to be directly related to ECM biomass and species composition in any of the tree species. Ectomycorrhizal species composition and number of mycorrhiza did not explain the CO(2) exchange results at different temperatures.     

Effects of ozone on growth and gas exchange of Eucalyptus globulus seedlings

To study the effects of a low concentration of ozone on growth and gas exchange in Eucalyptus globulus Labill. seedlings, ozone was applied for 37 days at a concentration of 50 ppb for 7 h daily under conditions of low light (250 micro mol m(-2) s(-1) (PAR)) and controlled temperature (20 degrees C). The seedlings exhibited extreme sensitivity to ozone. The ozone treatment reduced total plant biomass but had no effect on the partitioning of assimilate. Photosynthesis, stomatal conductance and internal CO(2) concentration were all reduced by ozone. The decline in photosynthesis was partly the result of direct effects of ozone on the stomata.     

Shoot and root sensitivity of containerized black spruce,white spruce and jack pine seedlings to late fall freezing

Damage to containerized forest seedlings due to freezing can occur in the fall or early winter in Canadian forest nurseries. The following spring, damage to shoots and impairment of growth is observed. The objectives of this experiment were to measure the impact of late fall low temperatures (0° to --30°C) on whole seedlings of the three most common species used for reforestation in Quebec: black spruce (BS), white spruce (WS) and jack pine (JP). Impacts of freezing temperatures on (i) whole seedling and apical bud mortality, (ii) shoot growth and root mortality, (iii) stem electrical resistance, (iv) shoot and root water relations, (v) concentrations of N, P, K, Ca, Mg, and total sugars in shoots were assessed. JP showed the highest rate of whole seedling mortality while WS showed the highest rate of apical bud mortality. JP was the most severely affected: destruction of the root system at low temperatures as well as a reduction of shoot growth and stem diameter and a decrease (more negative) in shoot and root water potential. WS showed a reduction of shoot growth despite no apparent damage to the root system at low temperatures. BS was not affected by temperatures as low as --30°C. Nutrient and sugar concentrations were not affected by low temperature treatments.     

Influence of foliar N on foliar soluble sugars and starch of red spruce saplings exposed to ambient and elevated ozone

Red spruce (Picea rubens Sarg.) trees growing at high elevation in the northeastern United States have experienced decline in recent years but seedlings have proved to be relatively tolerant of a wide range of environmental stresses in controlled studies. One possible reason for the wide tolerance to stress in seedlings is their inherently large pool of carbohydrate reserves, which is available for maintenance during and regrowth after periods of stress. We tested for the effects of foliar N and exposure to ozone on foliar carbohydrate reserves of 20-year-old naturally regenerated saplings. The trees were maintained in native soil in 360-l containers for 5 years before the experiment. The year before the experiment, trees were fertilized with N,P,K to provide a population of trees from N deficient to N sufficient. As foliar N decreased below 0.9%, length of current-year shoots and specific needle area of current-year needles declined. Foliar N concentration was correlated with foliar sugar and starch concentrations, but relationships varied with time of year. Before bud break, foliar carbohydrates and N, in general, were positively correlated, and date of bud break was delayed in N-deficient trees. During active growth, foliar soluble sugars and N were positively correlated, but starch concentrations were negatively correlated with N. By late September, neither starch nor sugar concentration was correlated with N concentration. Ozone and foliar N concentrations did not interact to change foliar carbohydrate concentrations or shoot and needle growth in this relatively short-term study.     

Leaf-level and whole-plant gas exchange characteristics of shortleaf pine exposed to ozone and simulated acid rain

Field-grown shortleaf pine (Pinus echinata Mill.) seedlings were exposed to ozone (O(3)) and simulated acid rain (SAR) in open-top chambers over three growing seasons. Ranges of O(3) and SAR spanned ambient levels found in the southern USA. Effects of O(3) on leaf-level and whole-plant gas exchange were characterized for a single measurement period immediately before the third summer of exposure. Decreased photosynthesis rates were attributed to O(3), but not SAR. Stomatal conductance decreased in response to O(3) exposure, and either increased or was unaffected by SAR. Increased internal CO(2) concentration (c(i)) in response to O(3) treatment indicated a greater effect of O(3) on photosynthetic capacity than stomatal conductance. Whole-seedling gas exchange characteristics indicated that whole-seedling carbon assimilation was more severely affected by O(3) than was evident from leaf-level gas exchange characteristics. Seedlings exposed to O(3) retained fewer flushes than seedlings grown in charcoal-filtered air.     

Atmospheric carbon dioxide concentration, nitrogen availability, temperature and the photosynthetic capacity of current-year Norway spruce shoots

Branches of field-grown Norway spruce (Picea abies (L.) Karst.) trees were exposed to either long-term ambient or to elevated CO2 concentrations ([CO2]) using the branch bag technique. The light-saturated photosynthetic rates (A(max)) of current-year shoots differing in nitrogen (N) status were measured at various temperatures and at either ambient (360 micromol mol(-1), AMB) or elevated (ambient + 350 micromol mol(-1), EL) [CO2]. The value of A(max) was determined at various intercellular [CO2]s (A/Ci curves) and used to normalize photosynthetic rates to the mean treatment C(i) values, which were 200 micromol mol(-1) (AMB) and 450 micromol mol(-1) (EL), respectively. Needle N status and temperature strongly affected A(max). The response to N increased with temperature, and the photosynthetic temperature optimum increased with N status. This was assumed to be a result of reduced mesophyll CO2 conductance. The relative increase of Amax in the EL treatment compared to the AMB treatment varied from 15 to 90%, and increased with temperature, but decreased with N status. Nevertheless, the absolute photosynthetic response to EL increased with shoot N status. The relative increase in the instantaneous response of A(max) to elevated [CO2] was about 20% higher than the long-term response, i.e., there was downward acclimation in Amax in response to elevated [CO2]. The photosynthetic temperature optimum increased 4 degrees C with either a short- or a long-term increase in [CO2]. The bag treatment itself increased A(max) by approximately 16% and the temperature optimum of A(max) by approximately 3 degrees C.     

Strain fields around knots in Norway spruce specimens exposed to tensile forces

Two-dimensional strain fields around knots in two Norway spruce specimens subjected to tension loading were detected using a contact-free measuring technique based on white-light digital image correlation. The first specimen included a traversing Edge knot, and the second one, a Centric knot. The development of strain fields as a function of load level was measured by consecutive cyclic load tests where one side of the specimen was studied during each test. The objectives were to examine to what extent the strain fields could be detected, to investigate the correlation between strain fields measured on different sides of a specimen and to analyse the strain distributions around the knots. The results show that the applied technique is useful for catching both overall and detailed information about the behaviour of knots in wood members exposed to loading. Clear wood defects that could not have been detected by neither visual inspection nor scanning were observed, and conclusions could be drawn regarding the release of internal stresses. The correlations between strain fields on different sides of the specimens were excellent, and the correspondence between measurement results and comparative finite element calculations was surprisingly good considering the fact that the employed FE models were fairly simple.     

Hebeloma inoculation on Norway spruce seedlings in solarized and infested soil

The effect of inoculation of Hebeloma sp. on the growth of Norway spruce (Picea abies) seedlings was studied, in a controlled environment, with soil samples naturally infested by Pythium spp., Fusarium spp. and Rhizoctonia solani, and with samples of soil in which the pathogens had been controlled by solarization. The best results were obtained in solarized soil, but mycorrhizae also enhanced plant growth in the presence of the whole soil-borne mycoflora. The combined use of inoeulation of a mycorrhizal fungus and solarization is discussed.     

Seasonal changes in chlorophyll fluorescence of white spruce seedlings from different latitudes in relation to gas exchange and winter storability

Variable chlorophyll fluorescence (F_var) was investigated as a tool in detection of distinct seasonal physiological changes in 1+0 intact white spruce seedlings. The loss of the characteristic F_var peak (Fp) between 0.8 and 1.0 s after illumination of dark adapted seedlings is an indication of regulation of photosynthetic activity in August. The peak represents excess photochemical water-splitting of photosystem II. We interpret its loss as a physiological indicator of the process of dormancy induction. Three dimensional (i.e. X[0-300 s], Y[rfu], Z[time of year] axis) F_var curves of non-stressed seedlings measured over 300 s followed a three phase change over the growing season. In actively growing seedlings, the portion of the Kautsky induction curve between 60 and 300 s was 0.4 relative fluorescence units (rfu) in northern (i.e. >56° latitude) seedlots and 0.6 in the southern (i.e. <56° latitude) range seedlot from August until early September. About mid-September curve features between 60 and 300 s decreased sharply to approximately zero (rhu) by October. Freeze test data indicated seedlings became frost hardy during this time. The third, or inactive phase was seen as flat line from 40 to 100 s. The portion of the curve after 100 s was responsive to short term temperature changes. White spruce seedlots of northern and southern B.C. latitudes having curve fluorescence peak (F_p) values at about 1 s of 0.6, and 0.8 (rhu) respectively, plus curve minimum (F_min) values at about 60 s which do not decrease further over a 2–3 week period represent stock which can safely be lifted for cold storage. The F_var attribute at 5 s after the actinic light is turned on (F_5s) correlates well with net photosynthesis (r² =0.61) during the growing season.     

Impact of soil pressure and compaction on tracheids in Norway spruce seedlings

We tested the effect of soil compaction on Norway spruce seedlings in terms of the size and theoretical volume flow rate of the tracheids. The results show that soil pressure limits growth in the diameter of the lumens of tracheids in all parts of seedlings studied. The tracheids of the roots with primary xylem had larger lumens than those of the roots and shoots with secondary xylem in both unloaded and loaded seedlings. This corresponds to the higher cumulative theoretical volume flow rate of the tracheids from roots with primary xylem than those from roots and shoots with secondary xylem. Although the volume flow rate of tracheids, according to the Hagen-Poiseuille law, was directly proportional to the quadratic power of the capillary diameter (tracheid lumen), the cumulative curve of the theoretical hydraulic volume flow rate was higher or relatively comparable in loaded seedlings. An explanation for these findings is that there were higher gradients of water potential values in roots and leaves in loaded seedlings because the lengths of the conductive pathways were 27% shorter than in unloaded seedlings. We hypothesise that trees have adapted to different stresses by shortening their conductive pathways to maintain a transpiration rate similar to that of non-stressed trees. These results concerning the impact of soil compaction on tracheid diameter and volume flow rate improve our understanding of the growth and functioning of different conifer organs and the mechanisms underlying the efficiency of water transport through the root xylem to the shoot.     

Effects of soil temperature on shoot and root growth and nutrient uptake of 5-year-old Norway spruce seedlings

Soil temperature is a main factor limiting root growth in the boreal forest. To simulate the possible soil-warming effect of future climate change, 5-year-old Norway spruce (Picea abies (L.) Karst.) seedlings were subjected to three simulated growing seasons in controlled environment rooms. The seedlings were acclimated to a soil temperature of 16 degrees C during the first (GS I) and third growing seasons (GS III), but were assigned to random soil-temperature treatments of 9, 13, 18 and 21 degrees C during the second growing season (GS II). In GS II, shoot diameter growth was lowest in the 21 degrees C treatment and root growth was lowest in the 9 degrees C treatment. In GS III, shoot height and root length growth improved in seedlings that had been kept at 9 degrees C during GS II, indicating compensatory growth in response to increased soil temperature. The temporary decrease in soil temperature had no long-lasting significant effect on seedling biomass or total nutrient uptake. At the end of GS III, fine roots of seedlings exposed to a soil temperature of 21 degrees C in GS II were distributed more evenly between the organic and mineral soil layers than roots of seedlings in the other treatments. During GS II and GS III, root growth started earlier than shoot growth, decreased during the rapid shoot elongation phase and increased again as shoot growth decreased.     

Carbon dioxide exchange in Norway spruce at the shoot, tree and ecosystem scale

Net CO2 exchange in a 35-year-old boreal Norway spruce (Picea abies (L.) Karst.) forest in northern Sweden was measured at the shoot (NSE), tree (NTE) and ecosystem levels (NEE) by means of shoot cuvettes, whole-tree chambers and the eddy covariance technique, respectively. We compared the dynamics of gross primary production (GPP) at the three levels during the course of a single week. The diurnal dynamics of GPP at each level were estimated by subtracting half-hourly or hourly model-estimated values of total respiration (excluding light-dependent respiration) from net CO(2) exchange. The relationship between temperature and total respiration at each level was derived from nighttime measurements of NSE, NTE and NEE over the course of 1 month. There was a strong linear relationship (r2 = 0.93) between the hourly estimates of GPP at the shoot and tree levels, but the correlation between shoot- and ecosystem-level GPP was weaker (r2 = 0.69). However, the correlation between shoot- and ecosystem-level GPP was improved (r2 = 0.88) if eddy covariance measurements were restricted to periods when friction velocity was > or = 0.5 m s(-1). Daily means were less dependent on friction velocity, giving an r2 value of 0.94 between shoot- and ecosystem-level GPP. The correlation between shoot and tree levels also increased when daily means were compared (r2 = 0.98). Most of the measured variation in carbon exchange rate among the shoot, tree and ecosystem levels was the result of periodic low coupling between vegetation and the atmosphere at the ecosystem level. The results validate the use of measurements at the shoot and tree level for analyzing the contribution of different compartments to net ecosystem CO2 exchange.     

Induction of callose in roots of Norway spruce seedlings after short-term exposure to aluminum

Callose (1,3-beta-glucan) is a suggested physiological indicator of aluminum (Al) toxicity in crop plants. It is not known if callose serves a similar function in forest trees, because quantitative data on callose formation in tree roots are limited, particularly under controlled conditions. To evaluate callose as a physiological indicator of Al toxicity in tree roots, we quantified callose formation in roots of Norway spruce (Picea abies (L.) Karst.) seedlings. Seedlings were grown in simulated soil solutions in the presence or absence (control) of Al under controlled conditions. In seedlings grown in solutions containing 280 microM Al, callose concentrations in roots were twice as high as control values after 6 h of Al treatment and 5 times higher than control values after 1 day. Thereafter, root callose concentrations gradually decreased and were only twice as high as control values after 7 days. The presence of various Al concentrations in the simulated soil solutions indicated that callose was induced by a relatively low Al concentration (84 microM). We conclude that callose in tree roots is an indicator of Al toxicity.