Anatomical and Histochemical Changes of Norway Spruce Buds Induced by Simulated Acid Rain

The study was focused on changes of anatomical and histochemical parameters of buds of 4-year-old Norway spruce (Picea abies L. Karst) trees subjected to simulated acid rain (SAR). Solutions of pH 2.9 and 3.9 were applied by spraying on shoot and/or by watering for two years. No macroscopic changes of buds or needles were observed in connection with SAR application and the only induced change was 2-week earlier onset of bud break in all treated variants compared to the control. Two-year treatment caused decrease in number of leaf primordia and increase in number of living bud scales in treated dormant buds while these parameters remained unchanged in the control buds. Treatments with solution of pH 2.9 caused decrease of flatness of bud apical meristem during the vegetative season. Increased activity of non-specific esterase in treated buds occurred during dormancy and bud break and the enhanced accumulation of phenolic compounds was detected at the beginning of shoot growth. Changes in histochemical parameters of bud tissues were induced mainly by spraying of shoots and can thus be qualified as primary damage.     

Comparison of Fine Root Dynamics in Scots Pine and Pedunculate Oak in Sandy Soil

In this study, we investigated the relationship between the seasonality of vegetation cover and that of fine root processes in a man-made forest in northern Belgium. Due to their contrasting foliar development, we expected different seasonal patterns of fine root growth and standing biomass between Pedunculate oak (Quercus robur L.), and Scots pine (Pinus sylvestris L.). Biomass and necromass of fine and small roots were estimated by repeated core sampling in February, April, June, August and October 2003. Measurements showed that Pedunculate oaks maintained more live fine roots in winter than Scots pines. However, Scots pines produced more than twice as much fine roots in spring, such that in summer both species had similar root mass. Scots pine root production started before-, but declined during leaf unfolding. Pedunculate oak roots, in contrast, started elongating only after bud break. For both species, fine root production peaked in JuneJuly, but was more than offset by drought-induced mortality at the end of July and early August. Summer drought in 2003 was exceptionally long and intense, significantly reducing leaf area, killing most new roots, and inhibiting root decomposition, such that the obtained results cannot be typical for this forest.     

Biodegradation of Trichloroacetic Acid in Norway Spruce/Soil System

Trichloroacetic acid (TCA) belongs to secondary atmospheric pollutants affecting the forest health. Distribution of [1,2-¹⁴C]TCA-residues and TCA biodegradation were investigated in 4-year-old nursery-grown trees of Norway spruce [Picea abies (L.) Karst.] in the whole plant/soil system. Radioactivity was monitored in needles, wood, roots and soil as well as in the air. During two weeks of exposure TCA was continuously degraded, especially in the soil. Estimates of radioactivity balance showed loss of radioactivity into the atmosphere in the form of ¹⁴CO₂; unincorporated [1,2-¹⁴C]TCA, chloroform, carbon monoxide and methane were not detected at all. TCA degradation to CO₂ was indicated also in the spruce needles. Moreover, it was found that soil litter contained [1,2-¹⁴C]TCA unavailable to microorganisms.     

pH buffering in acidic soils developed under Picea abies and Quercus robur – effects of soil organic matter, adsorbed cations and soil solution ionic strength

Soil solution chemistry, soil acidity andcomposition of adsorbed cations were determinedin two soil profiles developed under a mixedspruce (Picea abies and Piceasitchensis) stand and in one soil profiledeveloped under an oak (Quercus robur)stand. Soils under spruce were classified asSpodosols and soils under oak were classifiedas Inceptisols. All profiles were developed inthe same parent material; a Saahlian sandy tillcontaining less than 2% clay. In the mineralsoil, the contribution from mineral surfaces tothe total cation-exchange capacity (CEC_t)was estimated to be less than 3%. Soilsolution pH and the percent base saturation ofCEC_t [%BS = 100 (2Ca + 2Mg + Na + K)CEC_t ^–1] were substantially lower inthe upper 35–40 cm of the two Spodosols, ascompared to the Inceptisol. The total amount ofsoil adsorbed base cations (BC) did not differamong the three profiles on an area basis downto 1 m soil depth. Thus, soil acidification ofCEC_t due to net losses of BC could notexplain differences in soil pH and %BS amongthe soil profiles. A weak acid analogue, takingthe pH-effect of metal complexation intoconsideration, combined with soil solutionionic strength as a covariate, could describeboth the pH variation by depth within soilprofiles and pH differences between theInceptisol and the two Spodosol profiles. Ourresults confirm and extend earlier findingsfrom O and E horizons of Spodosols that theextent to which organic acid groups react withAl minerals to form Al-SOM complexes is a majorpH-buffering process in acidic forest soils. Wesuggest that an increasing Al-saturation of SOMis the major reason for the widely observed pHincrease by depth in acidic forest soils with apH less than approximately 4.5. Our resultsstrongly imply that changes in mass of SOM, theionic strength in soil solution and therelative composition of soil adsorbed Al and Hneed to be considered when the causality behindchanges in pH and base saturation isinvestigated.     

Image Analysis — Tool for Quantification of Histochemical Detections of Phenolic Compounds, Lignin and Peroxidases in Needles of Norway Spruce

Image analysis has become a powerful tool for the quantification of histochemical detections mainly in animal and medical sciences. There is not, however, much information about the accuracy of image analysis quantification of histochemical products and their comparison to biochemical analyses. Our study focused on the quantification of histochemically detected amounts of phenolic compounds, lignin and activity of peroxidases in the two youngest age classes of Norway spruce (Picea abies (L.) Karst) needles with the use of image analysis and its comparison with biochemical assays. The image analysis-determined amount of lignin and peroxidase activity was well correlated with its biochemical assay. On the other hand, no optical parameters of the reaction product of the histochemical reaction of phenolics correlated with the biochemical quantification. This proves that results of quantification of histochemical detection with image analysis greatly depend on the character and procedure of the histochemical proof. If it is done in full respect of the conditions of a histochemical reaction it can gain reliable and interesting results. Great advantage of the image analysis approach is quantification of the studied substance in different tissues within single sample as well as among different samples, which in many cases cannot be detected with biochemical tools. This revised version was published online in July 2006 with corrections to the Cover Date.     

Photosynthesis and transpiration of healthy and diseased spruce trees in the course of three vegetation periods

Summary CO₂- and H₂O-gas exchange of 20- to 25-year-old spruce trees from a plantation in the Hunsrück mountains were investigated over a period of 3 years. All measurements were made as pair comparisons, i.e., in each case the gas exchange of a damaged tree and of a relatively healthy tree in its immediate vicinity was measured simultaneously. A second plantation in the Westerwald mountains consisted of 18-year-old apparently healthy spruce trees. Pair comparison at this location meant comparison of two healthylooking trees. The investigations at both locations included diurnal course measurements of photosynthesis and transpiration, and light saturation curves and CO₂-saturation curves of photosynthesis. The reduced photosynthesis parameters of the phenotypically damaged trees at the Hunsrück location indicates massive damage to the photosynthetic apparatus. Measurements of H₂O-gas exchange showed that there are disturbances in stomatal regulation of the needles of damaged trees. As a result, the water use efficiency of these needles proved to be significantly lower. In addition, apparent photorespiration of the damaged trees was decreased, whereas their light- and CO₂-compensation points and their dark respiration were increased. In contrast to the Hunsrück plantation, no such effects were detectable when the healthy-looking Westerwald trees were subjected to pair comparison of gas exchange. Reduced photosynthetic capacity and disturbances of the stomatal regulation of the phenotypically damaged Hunsrück trees may be due to damage in the cellular membranes. Furthermore, a comparison of three growing seasons led to the conclusion that the gas exchange of spruce trees in their natural habitat is markedly influenced by climatic conditions.     

The Mitotic Activity of Norway Spruce Polyembryonic Culture Oscillates During the Synodic Lunar Cycle

The present paper tests the hypothesis, that a periodic fluctuation of mitotic activity of the embryonal tissue of the Norway spruce (Picea abies (L.) Karst.) is synchronnous with synodic lunar cycling. The increased mitotic index (MI) was observed under full moon, and the decreased MI around the first and third quarter. This revised version was published online in July 2006 with corrections to the Cover Date.     

Differences in Anatomical Structure and Lignin Content of Roots of pedunculate Oak and Wild Cherry-Tree Plantlets During Acclimation

The lignin contents and anatomical structure of roots of wild cherry (Prunus avium L.) and pedunculate oak (Quercus robur L.) plantlets were compared to explain differences in response during transfer from in vitro to ex vitro conditions. Lignification of cell walls increased significantly in both oak and cherry roots during the period of acclimation and finally lignin content of root tissues of in vitro propagated plantlets reached the levels not significantly different from seedlings grown in soil. Later on when secondary tissues appeared, lignified secondary xylem constituted most of the tissues of both species. The most conspicuous interspecific difference in root structure was the presence of phi-thickenings in cortical layers just outer to endodermis in cherry roots cultivated ex vitro. Formation of phi-thickenings was avoided in vitro and their presence thus seems to be under environmental control. Suberised well established exodermis was present in roots of oak but not detected in those of cherry. Very early development of exodermis in oak roots, preceding suberisation of endodermis, was recorded in vitro but not in well aerated soil. While multilayered and well-developed cork occurred in oak, only thin walled and less suberised secondary dermal tissues were found in cherry.     

Different Behaviour of Indole-3-Acetic Acid and Indole-3-Butyric Acid in Stimulating Lateral Root Development in Rice (Oryza sativa L.)

The plant hormone auxin has been shown to be involved in lateral root development and application of auxins, indole-3-acetic acid (IAA) and indole-3-butyric acid (IBA), increases the number of lateral roots in several plants. We found that the effects of two auxins on lateral root development in the indica rice (Oryza sativa L. cv. IR8) were totally different from each other depending on the application method. When the roots were incubated with an auxin solution, IAA inhibited lateral root development, while IBA was stimulatory. In contrast, when auxin was applied to the shoot, IAA promoted lateral root formation, while IBA did not. The transport of [³H]IAA from shoot to root occurred efficiently (% transported compared to supplied) but that of [³H]IBA did not, which is consistent with the stimulatory effect of IAA on lateral root production when applied to the shoot. The auxin action of IBA has been suggested to be due to its conversion to IAA. However, in rice IAA competitively inhibited the stimulatory effect of IBA on lateral root formation when they were applied to the incubation solution, suggesting that the stimulatory effect of IBA on lateral root development is not through its conversion to IAA.     

Acclimation of crown structure to drought in Quercus robur L.—intra- and inter-annual variation of abscission and traits of shed twigs

In pedunculate oak (Quercus robur L.) the architecture of the crown is strongly influenced by age and vigour of the tree. In old oak trees cladoptosis is a major mechanism on the transformation of crown architecture. Although it can be seen quite regularly, the causes and timing of shedding of twigs as well as the quantity of affected branches remain unclear. Because abscission is often used as an indicator of reduced vigour or stress in the assessment of stand and ornamental tree health, it deserves detailed investigation, especially in the context of oak decline. We studied the inter- and intra-annual variation of abscission in six stands across the eastern part of Germany in order to identify possible triggering events and controlling factors of abscission.On average, the number of twigs abscised per year increased from 1999 to 2001. While in 1999 approximately 100 abscised twigs per m² per year were shed, this number increased to 250 per m² in 2001. The majority of twigs was actively shed, a significant proportion of the remaining twigs was partly abscised. Only a small part of the abscised twigs had leaves attached to them. From June to September 20 per cent of the twigs had leaves, in the remaining months of the year less than 10 per cent.The analysis of almost 30,000 twig fragments over the course of 3 years demonstrated at least partial control by the tree of the process of abscission. The loss of the terminal bud and the formation of a male flower are traits correlated with abscission.Our data do not support the widely held belief that cladoptosis is mainly an immediate reaction to drought stress that reduces transpiring leaf area. The proportion of leafy twigs was quite low and the main peak of abscission occurred in late autumn, thus having little immediate effect on transpiration. In addition, we observed a time lag of 3 weeks between the onset of drought stress and increased levels of abscission.     

Control Mechanisms of Photosynthetic Capacity Under Elevated CO2 Concentration: Evidence from Three Experiments with Norway Spruce Trees

Twelve-year-old Norway spruce (Picea abies [L.] Karst.) were exposed to ambient (AC) or elevated (EC) [ambient + 350 mol(CO₂) mol^–1] CO₂ concentration [CO₂] using the facilities of open-top-chambers (OTCs) and glass domes (GDs). A combination of gas exchange measurements and application of a biochemical model of photosynthesis were used for the evaluation of CO₂ assimilation characteristics. Morphological change was assessed on the base of specific leaf area (SLA). Nitrogen (N) content in the assimilation apparatus was considered a main factor influencing the biochemical capacity. Three experiments confirm the hypothesis that an adjustment of photosynthetic capacity under EC is controlled by the combination of biochemical, morphological, and physiological feedback mechanisms. We observed periodicity of down-regulation of photosynthetic capacity (Experiment No. 1) during the vegetation seasons. In the spring months (May–June), i.e. during the occurrence of active carbon sink associated with the formation of new foliage, up-regulation (10–35 %) of photosynthetic capacity (P _Nsat) was observed. On the contrary, in the autumn months (September–October) down-regulation (25–35 %) of P _Nsat was recorded that was mainly associated with reduced carbon sink strength and biochemical change, i.e. decrease of N status (up to 32 %) and accumulation of saccharides (up to 72 %) in leaves. Different adjustments of photosynthetic activities were observed in current (C) and one-year-old (C-1) needles exposed to EC (Experiment No. 2). Strong down-regulation of P _Nsat and the diminution of the initial stimulation of photosynthetic rate (P _Nmax) was associated with decreases of both ribulose-1,5-bisphosphate carboxylase/oxygenase carboxylation activity (by 32 %) and RuBP regeneration (by 40 %). This performance was tightly correlated with the absence of active carbon sinks, decrease of N content, and starch accumulation in C-1 needles. Finally, different responses of sun- and shade-adapted needles to EC (Experiment No. 3) were associated with the balance between morphological and biochemical changes. Observed P _Nsat down-regulation (by 22 %) of exposed needles in EC was predominantly caused by effects of both higher assimilate accumulation and stronger N dilution, resulting from higher absolute photosynthetic rates and incident irradiances in the upper canopy.     

Chlorophyll a Fluorescence Response of Norway Spruce Needles to the Long-Term Effect of Elevated CO2 in Relation to Their Position Within the Canopy

The long-term impact of elevated CO₂ concentration on photosynthetic activity of sun-exposed (E) versus shaded (S) foliage was investigated in a Picea abies stand (age 12 years) after three years of cultivation in adjustable-lamella-domes (ALD). One ALD is supplied with either ambient air [ca. 350 µmol(CO₂) mol^–1; AC-variant) and the second with elevated CO₂ concentration [ambient plus 350 µmol(CO₂) mol^–1; EC-variant). The pronounced vertical profile of the photosynthetically active radiation (PAR) led to the typical differentiation of the photosynthetic apparatus between the S- and E-needles in the AC-variant estimated from the irradiance-responses of various parameters of the room temperature chlorophyll (Chl) a fluorescence parameters. Namely, electron transport rate (ETR), photochemical efficiency of photosystem 2, PS2 (_PS2), irradiance-saturated values of non-photochemical quenching of minimum (SV₀) and maximum (NPQ) fluorescence levels, and photochemical fluorescence quenching (q_p) at higher irradiances were all significantly higher for E-needles as compared with the S-ones. The prolonged exposure to EC did not cause any stimulation of ETR for the E-needles but a strongly positive effect of EC on ETR was observed for the S-needles resulting in more than doubled ETR capacity in comparison with S-needles from the AC-variant. For the E-needles in EC-variant a slightly steeper reduction of the _PS2 and q_p occurred with the increasing irradiance as compared to the E-needles of AC-variant. On the contrary, the S-needles in EC variant revealed a significantly greater capacity to maintain a high _PS2 at irradiances lower than 200 µmol m^–2 s^–1 and to prevent the over-reduction of the PS2 reaction centres. Moreover, compared to the AC-variant the relation between SV₀ and NPQ exhibited a strong decrease (up to 72 %) of the SV₀-NPQ slope for the E-needles and an increase (up to 76 %) of this value for the S-needles. Hence the E- and S-foliage responded differently to the long-term impact of EC. Moreover, this exposure was responsible for the smoothing of the PAR utilisation vertical gradient in PS2 photochemical and non-photochemical reactions within the canopy.     

Changes in the content and constituents of the cuticular wax of Picea abies (L.) Karst. in relation to needle ageing and tree decline in five European forest areas

Cuticular wax composition of healthy and and declining mature Norway spruce trees [Picea abies (L.) Karst.] was investigated in five European forest areas. The amount of extracted wax and the content of alkanes and secondary alcohols were analysed as a function of the factors sample area (five areas, detailed below), needle age (current year to 2 years) and decline class (Class O to Class 2). Using a GC-MS, alkanes from C₂₀ to C₃₁ and the following alcohols were quantified: 10-nonacosanol, 5,10-nonacosandiol, 4,10-nonacosandiol and the triterpenol 24-methylenecycloartanol. According to our results, the total wax content as well as the alkane and alcohol content of waxes show a large variation corresponding to sample area and needle age. Ageing caused a highly significant increase in alkane content and a highly significant decrease in total wax and alcohol content. The decline class significantly influenced only the content of the long chain alkane C₃₁ (increase), the secondary alcohol 10-nonacosanol (decrease), and the triterpenic alcohol (decrease). Total wax weight was not influenced by tree damage. Thus, according to our results, needle ageing and progressive tree damage are correlated to different changes in the examined parameters.     

Nitrogen mineralization in high elevation forests of the Appalachians. I. Regional patterns in southern spruce-fir forests

Annual and seasonal rates of net nitrogen mineralization were determined for 19 sites in the spruce-fir forests of the Southern Appalachian Mountains. These sites included high and low elevation stands of red spruce (Picea rubens Sarg.) and Fraser fir (Abies fraseri (Pursh.) Poir.) on east and west exposures on Whitetop Mountain, Virginia; Mt. Mitchell, North Carolina; and Clingman's Dome in the Great Smoky Mountains National Park. Mineralization rates were determined using in situ soil incubations in PVC tubes with ion exchange resin bags placed in the bottom of the tubes to collect leachate. Throughfall was collected in resin bags placed in the top of the tubes. Average initial NH₄-N + NO₃-N ranged from 0.6 to 4.8 kg N/ha across all plots, and average mineralization rates ranged from 26 to 180 kg-N ha⁻¹ yr⁻¹. Throughfall ranged from 18 to 32 kg-N ha⁻¹ yr⁻¹ with NH₄-N accounting for about two-thirds of the throughfall N across all sites. Throughfall and mineralization rates were not related to elevation or exposure. The high rates of N mineralization and relatively high nitrate concentrations indicate that leaching losses of nitrogen and associated cations could be substantial. Requests for offprints     

Polar Transport of Indole-3-Acetic Acid in Relation to Rooting in Carnation Cuttings: Influence of Cold Storage Duration and Cultivar

The influence of cold storage of cuttings on the transport and metabolism of indole-3-acetic acid (IAA) and the rooting were studied in two carnation (Dianthus caryophyllus L.) cultivars (Oriana and Elsy), which are known to exhibit very distinct rooting characteristics. The percentage of rooting at 11 d after planting increased with the storage period particularly in Oriana, but the values in Elsy were higher than in Oriana. Auxin transport was measured by applying ³H-IAA to stem sections. Irrespective of the section localization, the oldest node (node) or the basal internode (base), the transport increased as the storage period increased from 2 to 12 weeks in Oriana and from 2 to 8 weeks in Elsy cuttings. The auxin transport rate was higher in bases than in nodes and also in Elsy than in Oriana at a given storage period. IAA oxidation and hydrolyzation of IAA conjugates (determined by extracting the sections with acetonitrile and NaOH once the basipetal IAA movement ceased after a 24 h transport period) showed a negative, highly significant correlation with the amount of IAA transported. Although the rooting percentage and IAA transport were higher in Elsy than in Oriana, the differences in rooting between the cultivars could not be explained solely by differences in IAA transport.     

Contents of ATP and ADP in needles of Norway spruce in relation to their development,age, and to symptoms of forest decline

Summary Pool sizes of ATP and ADP were analysed in freeze-stopped, lyophilised homogenates of needles from Norway spruce [Picea abies (L.) Karst.]. Control experiments in which possible changes in adenylate pools during sample acquisition were investigated did not reveal significant differences between needles taken from branches in situ or within a 30-min period after cutting off a branch. In addition, pool sizes of ADP and ATP were not affected by changes in light intensity (between 60 and 1500 E^*-m^-2*s^-1), which inevitably occur when samples have to be taken from the upper region of older trees. Levels of ATP and ADP showed considerable seasonal changes (May through October) with the highest ratios of ATP/ADP in developing needles. In general, there was a tendency towards increased ratios of ATP/ADP with increasing needle age. This observation was corroborated by analyses of needles from spruce trees of different age and growing under different conditions. Needles from declining trees or from trees specifically fumigated with low concentrations of ozone and sulphur dioxide had significantly increased ratios of ATP/ADP compared to controls. The results are discussed with respect to physiological responses connected with natural senescence and induced ageing.     

Fine Root Production and Turnover in a Norway Spruce Stand in Northern Sweden: Effects of Nitrogen and Water Manipulation

Fine root length production, biomass production, and turnover in forest floor and mineral soil (0–30 cm) layers were studied in relation to irrigated (I) and irrigated-fertilized (IL) treatments in a Norway spruce stand in northern Sweden over a 2-year period. Fine roots (<1 mm) of both spruce and understory vegetation were studied. Minirhizotrons were used to estimate fine root length production and turnover, and soil cores were used to estimate standing biomass. Turnover was estimated as both the inverse of root longevity (RTL) and the ratio of annual root length production to observed root length (RTR). RTR values of spruce roots in the forest floor in I and IL plots were 0.6 and 0.5 y⁻¹, respectively, whereas the corresponding values for RTL were 0.8 and 0.9 y⁻¹. In mineral soil, corresponding values for I, IL, and control (C) plots were 1.2, 1.2, and 0.9 y⁻¹ (RTR) and 0.9, 1.1, and 1 y⁻¹ (RTL). RTR and RTL values of understory vegetation roots were 1 and 1.1 y⁻¹, respectively. Spruce root length production in both the forest floor and the mineral soil in I plots was higher than in IL plots. The IL-treated plots gave the highest estimates of spruce fine root biomass production in the forest floor, but, for the mineral soil, the estimates obtained for the I plots were the highest. The understory vegetation fine root production in the I and IL plots was similar for both the forest floor and the mineral soil and higher (for both layers) than in C plots. Nitrogen (N) turnover in the forest floor and mineral soil layers (summed) via spruce roots in IL, I, and C plots amounted to 2.4, 2.1, and 1.3 g N m⁻² y⁻¹, and the corresponding values for field vegetation roots were 0.6, 0.5, and 0.3 g N m⁻² y⁻¹. It was concluded that fertilization increases standing root biomass, root production, and N turnover of spruce roots in both the forest floor and mineral soil. Data on understory vegetation roots are required for estimating carbon budgets in model studies.     

Biochemical symptoms of stress in the mycorrhizal roots of Norway spruce (Picea abies)

Summary The mycorrhizal activity of spruce in a mixed-wood forest was monitored over 1 year by measuring biochemical characters in fine roots of six canopy trees and of a regrowth stand. The concentration of adenosine 5-triphosphate (ATP), a measure of living biomass, showed two peaks per year, one at bud break and one after main shoot growth. The concentration of storage polysaccharides in mycorrhizae showed the same cycles even more pronouncedly. It is proposed that these changes reflect growth and senescence of mycorrhizae and that the timing of the cycles is controlled by translocation of assimilates from the shoot. Differences between mycorrhizae collected from canopy trees and the regrowth stand were small and not significant. Characters known to be related to fungal activity of the mycorrhizal symbiosis (concentration of trehalose, glucose uptake, respiration) also varied little among the six canopy trees. Large differences among fine-root samples from different canopy trees, however, were detected in the concentrations of ATP and storage polysaccharides, measures which seemed to be physiologically integrated within trees. If low concentrations in roots precede losses of foliage from trees, these two symptoms could be used as early indicators of growth decline in individual spruce trees.     

Photosynthetic capacity,chloroplast pigments,and mineral content of the previous year's spruce needles with and without the new flush: analysis of the forest-decline phenomenon of needle bleaching

Summary Spruce (Picea abies) damage in the Fichtelgebirge (FRG) occurs as needle bleaching and a depression of CO₂ assimilation. Such injury may primarily result from the direct, above-ground effects of air pollution or indirect, below-ground changes in mineral uptake.Typically, the new flush of spruce needles is green and exhibits high photosynthetic capacity. Mies and Zöttl concluded that the older foliage is damaged when nutrients are withdrawn to supply the current year's needles. By removing the terminal buds of single branches in the spring, we produced an experimental set of the previous year's needles with greater mineral reserves than the control needles. During the course of the growing period, the performance of the experimental needles, which lacked competition from the new flush, was compared to that of the control needles of the same age-class on intact branches with the new flush.Throughout the experiment, chloroplast pigments of a healthy control tree were not affected by the elimination of the new flush. However, the chlorophyll and carotenoid content as well as the photosynthetic capacity of the previous year's needles on those branches of a heavily damaged tree where the new flush had been eliminated increased substantially. This increase was associated with an increase in minerals, which seemed to be deficient in the control needles with the new flush. Thus, in contrast to needles of the same age-class on intact branches with undisturbed new growth in the same atmospheric environment, the experimental needles escaped bleaching and a decrease in photosynthesis. It would seem that the bleaching and the loss in photosynthetic capacity typical of trees damaged by forest decline indirectly result from nutrient deficiencies through soil environment changes and/or root damage than directly from atmospheric pollutants.     

Modeling Response of N Addition on C and N Allocation in Scandinavian Norway Spruce Stands

Because nitrogen is considered to be the major growth-limiting element in boreal forests, the increasing nitrogen availability from deposition should lead to increasing growth. We have tested this assumption by simulating, with a simple model, carbon and nitrogen development in seven long-term fertilization experiments in three Nordic countries. The only differences between sites in the model are climate, the ambient nitrogen deposition, nitrogen fertilization regimes, and the initial conditions at the start of the experiment. The model simulates the observed stand development well as long as nitrogen remains the limiting factor. The simulated retention of deposited nitrogen is in general low (less than 50%), whereas retention of fertilizer nitrogen is higher. This seems to imply that the higher production in the fertilized stands will not be maintained once the fertilization is stopped. The model results also indicate that the major effect of climate on site productivity is through soil processes, not tree physiology.