Does timing of boron application affect needle and bud structure in Scots pine and Norway spruce seedlings?

Loss of apical dominance is a well-known boron (B) deficiency symptom in trees. Recent field studies indicate that B deficiency may cause irreversible damage in emerging leader buds leading to bushy growth, and changes in developing needles in mature Norway spruce trees. We experimentally studied if timing of B application affects needles and buds of Scots pine (Pinus sylvestris) and Norway spruce (Picea abies) seedlings with low initial B levels. The treatments were: no B (B0); B supply from the beginning of the simulated summer (B1); starting soon after bud burst (B2) and starting at the occurrence of first needle primordia in new spruce buds (B3). At the end of the experiment, B concentration in B1 was 23 mg kg⁻¹ (pine) or 17 mg kg⁻¹ (spruce) and lower in the later applications. In B0 it was at deficiency limit. In B0, B2 and B3, there were fewer sclerenchyma cells, and cavities occurred in vascular cylinders in pine needles, and in spruce buds there were more tanniferous cells in the primordial shoots compared to B1. Furthermore, in all but B1 there was cell collapse in the bud apex of some spruce seedlings. The experimentally induced changes were the same as earlier reported in B deficient conifers in the field, and indicate, similarly as in the field that adequate B is necessary throughout the growing season for healthy growth, particularly for spruce. The differences between spruce and pines are due, at least partly, to the differences in time frame of needle development and in the differences in development of conducting tissues in the buds.     

Does ice crystal formation in buds explain growth disturbances in boron-deficient Norway spruce?

Loss of apical dominance in boron-deficient trees has been suggested to be due to frost damage of terminal buds and leaders. Excessive nitrogen (N) supply can exacerbate boron (B) deficiency by the dilution-effect. N may also have direct effects on winter hardiness. We studied frost hardening of buds of Norway spruce (Picea abies L. Karst.) in healthy-looking trees and in trees with growth disturbances. The effect of B and N on frost hardiness was studied in a factorial fertilisation experiment during cold acclimation. Frost hardiness was determined by differential temperature analysis (DTA) and scoring of visual damage. In a DTA profile of apical buds with a piece of stem, low-temperature exotherm (LTE) predicted bud injury, while two of the observed high-temperature exotherms and two of the observed intermediate-temperature exotherms were non injurious. Appearance of LTE followed changes in air temperature. The risk of frost damage was not affected by fertilisation treatments or previously observed growth disturbances. However, when the bud structure was deformed by severe B deficiency, the supercooling ability disappeared. Such buds are probably killed by freezing in nature and therefore, frost damage may play a secondary role in the development of growth disturbances.     

Cold acclimation was partially impaired in boron deficient Norway spruce seedlings

Susceptibility of trees to freezing injury has been suggested to increase in boron (B) deficiency but there is no experimental evidence to support this proposition. In this study, Norway spruce (Picea abies L. Karst.) seedlings were cultivated for two growing seasons in deficient, intermediate and ‘optimal’ B levels. Cold hardening of the seedlings was measured after the second growing season. Freezing tolerance in tips of shoots, needles, stems and roots was determined by controlled freezing tests and electrolyte leakage method, and that of buds, in addition, by differential thermal analysis (DTA). Electrical impedance was used to monitor changes in the apoplastic space during cold acclimation. Root dry weight and shoot height growth were lower in B deficiency. Cold acclimation of buds and stems was reduced by B deficiency. When hardened seedlings were subjected to subzero temperatures for 3 weeks, extracellular electrical resistance of stems became the highest at the lowest B supply which was probably due to decreased desiccation tolerance. As a conclusion, susceptibility to freezing damage may be increased by B deficiency in Norway spruce trees.     

Evidence for the involvement of biotic factors in the yellowing disease of Norway spruce (Picea abies [L.] Karst.) at higher elevations of the Bavarian Forest

Extensive investigations on the fine root status of declining and healthy spruce were conducted in several stands at higher elevations of the Bavarian Forest heavily affected by needle yellowing. In most of the root parameters recorded, yellowing trees had significantly lower values than neighbouring green trees. Tight correlations were found between decreasing fine root density and crown transparency, degree of yellowing (increasing) and needle Mg (Ca) contents (decreasing), respectively. Although growing on the same substrate, green trees showed much better Mg (Ca) nutrition than yellow trees, indicating that poor fine root status contributes to Mg (Ca) deficiency in yellowing spruce. Experiments with spruce seedlings growing in soil samples from yellowing stands proved that needle symptoms can easily be reproduced on the seedlings under controlled conditions (i.e. in the absence of adverse atmospheric factors). Furthermore, reduced fine root systems and severe root damage were observed on seedlings grown in soils from yellowing stands, but not on those in soils from green stands. Adding a layer of soil from a yellow stand to a soil from a green stand caused a decrease of root parameters. Needle as well as root symptoms in these experiments were largely ameliorated after soils had been heat (autoclaving, sterilisation) or fungicide treated. Plants from treated soils had significantly longer roots and more root tips. The results of our study indicate that Mg deficiency leading to severe needle yellowing in stands at higher elevations of the Bavarian Forest is at least partially mediated by fine root disorders. Also, strong evidence is presented that fine root damage on trees in the affected stands is caused by soilborne micro-organisms, most likely fungi. Their exact identity, however, still remains to be unravelled.     

An analysis of needle yellowing in healthy and chlorotic Norway spruce(Picea abies) in a forest decline area of the Fichtelgebirge(N.E. Bavaria)

Summary In a forest decline area (Fichtelgebirge, N.E. Bavaria, FRG), annual time courses of chloroplast pigments in both healthy and chlorotic Norway spruce were studied. The seasonal time courses of green and apparently healthy trees did not generally differ from those reported in the literature for spruce trees of other regions. Chlorophyll content increased from May to October, remained relatively constant or declined slightly during the fall and early winter, and finally decreased markedly from March to early May when pigment is at its minimum before bud break. The annual maximal chlorophyll content increased with needle age from the current year's needles to 4-yearold needles. While carotene content reached its highest concentration in August, the xanthophylls did not peak until February or March. Pigment dynamics of chlorotic trees with lower concentrations, corresponded to those of undamaged trees. Chlorophyll deficits resulted from less pigment formation as well as pigment loss during the growing period. Even when the content of total chlorophyll was low, the ratio of chlorophyll a/chlorophyll b remained almost unchanged. In conjunction with the chlorophyll reductions, a decrease in the chlorophyll/carotenoid ratio and an increase in the xanthophyll/carotene ratio occurred. The periods of needle-chlorophyll reduction did not correlate with those periods of highest concentrations of atmospheric sulphur dioxide, the main air pollutant at the stand. However, chlorophyll formation ceased in the older needle age classes of chlorotic trees when the new flush was sprouting, indicating that nutritional deficiencies affect needle yellowing more than possible direct needle damage by air pollutants.     

Seasonal variation in the monoterpenes in needles of Picea abies (L.) Karst.

Summary This investigation was conducted to obtain information about the fluctuations in composition and amount of needle monoterpenes during the development of spruce needles. Studies conducted with two Norway spruce clones clearly revealed the existence of fluctuations. In juvenile needles, the amounts of the oxygenated terpenes increase constantly with age during the first 2 months of needle growth. The hydrocarbon terpenes dominate within the first weeks, some of them even showing a very distinct first maximum within the first 3 weeks after bud burst. All terpenes, including the oxygenated ones, have a maximum in June/July, which favours the hypothesis of a substitution of the hydrocarbons later on. There are significant changes even in mature needles of Norway spruce. The terpene level of 1-year-old needles of the clonal trees increased from spring to early summer and then dropped again towards winter. In addition, fluctuations in mature needles were shown for a set of ten wild trees. Needles of the same age class, which emerged in 1986, were sampled 4 times from 1986 to 1988. The needle terpene concentrations of the 1 -year-old needles were considerably lower in spring at the time of bud burst than in autumn. The terpene level of older needles thus seems to be influenced by biosynthetic and catabolic activities.     

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.     

Performance of two Picea abies (L.) Karst. stands at different stages of decline

Summary This is the first in a series of papers on the growth, photosynthetic rate, water and nutrient relations, root distribution and mycorrhizal frequency of two Norway spruce forests at different stages of decline. One of the stands was composed of green trees only while the other included trees ranging in appearance from full green crowns to thin crowns with yellow needles. In this paper we compare the growth and carbohydrate relations of the two stands and examine relationships among growth variables in ten plots. The declining stand produced 65 percent of the wood per ground area compared with the stand in which all trees were green because its foliage produced less wood at any level of leaf area index. The difference in foliage efficiency between the sites could not be explained by differeneces in climate, competition or stand structure. The declining stand appeared to have lower carbon gain as indicated by a smaller increase in reserve carbohydrates before bud break, and weaker sinks for carbohydrates as indicated by less use of the stored carbohydrates than the healthy stand. Thus, growth reduction was probably related to factors which affect both photosynthesis and, even more, the sinks for carbohydrate.     

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.     

Photosynthetic performance,chloroplast pigments,and mineral content of various needle age classes of spruce (Picea abies) with and without the new flush: an experimental approach for analysing forest decline phenomena

Summary Damage in the older needles of Norway spruce [Picea abies (L.) Karst.] in the Fichtelgebirge (NE Bavaria, FRG) appears to result primarily from nutrient imbalances rather than from direct effects of air pollutants on the mesophyll of the needles. Support for this conclusion was obtained by altering the nutrition of older needles through the removal of terminal buds on several branches from a damaged and an undamaged spruce tree in spring. Various photosynthetic parameters, as well as the chloroplast pigment and nutrient concentrations, of 1- to 3-year-old needles on manipulated branches were compared with those of branches on which the new flush was allowed to develop during the course of the growing period. Removal of terminal buds affected only the 1-year-old needles. Elimination of the new flush resulted in a higher Ca and Mn content of the needles of the undamaged tree. This treatment also resulted in an increase of the photosynthetic capacity (under saturating light and CO₂ conditions), carboxylation and light use efficiency, as well as net photosynthesis under natural conditions of the 1-year-old needles on the yellow chlorotic tree. This was accompanied by higher chlorophyll concentrations and an increase in Mg, Ca, Mn, and Zn content, and no visible signs of chlorosis developed in the experiment. By contrast, the needles of twigs in which the new flush was allowed to develop exhibited reductions in mineral content in the middle of the year. This was especially true for the elements Mg and Ca, and was accompanied by needle chlorosis and a depression of the capacity of photosynthesis. Thus it appears that there is a close relationship between the development of needle damage and nutrient imbalances in spruce. The retranslocation of elements from the 1-year-old needles to the new flush seems to play a major role in the development of needle bleaching. This approach thus supports the hypothesis described above and confirms a preliminary test with a similar experimental design, which had been conducted earlier.     

Herbivory modifies conifer phenology: induced amelioration by a specialist folivore

Herbivory by Zeiraphera canadensis Mut. & Free. (Lepidoptera: Tortricidae), an early season folivore of white spruce [ Picea glauca(Moench) Voss], has been associated with a shift in the timing of bud burst by its host during the subsequent year. We tested the hypothesis that a herbivory-induced shift in the phenology of bud development improves the window for colonisation of white spruce buds by Z. canadensis. Feeding on cortical tissue of elongating shoots caused the destruction of apical buds and an interruption of apical dominance in the year following herbivory. White spruce compensated for damage with the activation of dormant buds; mainly at proximal positions along shoots. As a result, half of all active buds on previously damaged branches were located immediately adjacent egg sites (i.e. previous year's bud scales), whereas <10% of active buds on intact shoots were situated there. More than 40% of newly emerged larvae colonised the basal buds of damaged shoots versus just 10% for intact shoots. Previous herbivory also influenced the initiation of bud burst. All buds flushed 2 days earlier on damaged shoots and date of bud burst was inversely correlated to bud density, indicating that short damaged shoots with large numbers of buds were stronger sinks for nutrients required for bud development. Egg hatch was best synchronized with early bursting buds on damaged branches. As a consequence, 89% of first-instar larvae successfully colonised buds on damaged branches while only 55% were successful on undamaged branches. Improved survival of larvae in the year following herbivory was a direct result of the evolved response by white spruce to the interruption of apical dominance. The pattern of herbivory by Z. canadensis may have evolved as a strategy to enhance the quality of white spruce for their offspring.     

受害马尾松针叶营养及次生物质含量与思茅松毛虫种群参数的相关分析

为了揭示受害马尾松Pinus massoniana针叶主要化学物质变化对思茅松毛虫Dendrolimus kikuchii Matsumura种群变动的影响, 本研究以受思茅松毛虫危害程度不同（轻度、 中度和重度）的马尾松针叶喂养思茅松毛虫幼虫, 测定思茅松毛虫饲养种群的特征参数和不同受害程度马尾松针叶营养物质与次生物质含量, 用综合相关系数分析法对试验结果进行分析。结果表明： （1）随植株受害程度加重, 针叶黄酮含量增加, 可溶性糖、 蛋白质、 多糖含量减少, 各龄幼虫平均历期延长、 死亡率升高。单宁和总酚含量的变化与各龄幼虫平均历期、 死亡率之间没有显著的相关性; （2）除6龄幼虫外, 其他各龄幼虫的平均历期、 死亡率均与松针营养物质和次生物质含量有直接的和综合的相关性; （3）随受害程度加重, 7龄幼虫体重、 幼虫平均取食量、 蛹重、 化蛹率、 雌性比、 每雌生殖力减小。单宁、 总酚含量的变化对7龄幼虫体重、 幼虫平均取食量、 蛹重、 幼虫平均死亡率、 化蛹率、 雌性比、 每雌生殖力都没有显著影响。总体上, 松针营养和次生物质含量对思茅松毛虫种群参数有重要影响, 其重要性依次为可溶性糖＞蛋白质＞多糖＞黄酮, 单宁和总酚的作用相对较小。     

Defoliation of Sitka spruce by the European spruce sawfly, Gilpinia hercyniae (Hartig): a retrospective analysis using the needle trace method

Abstract 1 During 1968–74, an outbreak of the European spruce sawfly Gilpinia hercyniae, defoliated many stands of Sitka spruce in commercial forests in mid‐Wales. The needle trace method was used to determine retrospectively the temporal pattern and intensity of defoliation in a stand of Sitka spruce in Hafren Forest that had been damaged severely at the time of the sawfly outbreak. 2 An initial calibration experiment, designed to test the reliability of the needle trace method when applied to Sitka spruce, indicated that artificially induced defoliation of up to 75% was detected by the technique with an accuracy of ±7%. Higher rates of defoliation were underestimated by up to 17%. 3 For the main sample of trees, the needle trace method demonstrated that retention of needle sets (average needle retention) was reduced by up to 33–38% over a 10‐year period coincident with the G. hercyniae outbreak and the years immediately afterward when the canopy was recovering. 4 Analysis of needle loss within separate needle cohorts and age‐classes revealed that 1‐year‐old needles were the most severely defoliated. The density of 1‐year‐old needles was reduced by 51–78% in 1970–73. 5 Defoliation at the time of the G. hercyniae outbreak was associated with reductions in annual height, radial, and volume increments of 24–49%, 30–59% and 32–56%, respectively. Radial and volume increments suffered their greatest reductions 1 year later than height increments. 6 The study demonstrates that the needle trace method can be applied successfully to Sitka spruce to quantify defoliation caused by an insect outbreak occurring many years previously, and that the technique can provide data on needle loss that is valuable for interpreting reductions in tree growth.     

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 incidence of mistletoe (<Emphasis Type="Italic">Viscum album</Emphasis> ssp. <Emphasis Type="Italic">abietis</Emphasis>) on silver fir (<Emphasis Type="Italic">Abies alba</Emphasis>) in Croatia

The research on incidence of mistletoe (Viscum album ssp. abietis) on silver fir (Abies alba) was carried out in natural fir stands in Croatia. In the area of Gorski Kotar 32.8 % of the examined dominant fir trees were infected with mistletoe. The mistletoe incidence was presented according to the damage degrees of silver fir from 0 (healthy trees) to 4 (dead trees), rated by the crown defoliation and needle discoloration. With the increase of incidence, mistletoe spreads more onto more vital, i.e. less damaged trees. In the Dinaric Alps 28.6% of the examined fir trees were infected, this percentage amounting to 27.1% for the mountainous regions between the Sava and Drava rivers. The site and stand parameters (exposure, elevation, site quality, forest community and crown closure) were analysed in order to establish whether there was a correlation between these parameters and mistletoe incidence. There was a negative correlation between the elevation and mistletoe incidence. Among the compartments with a closed stands there were considerably less compartments with higher mistletoe incidence than among compartments with a sparsely closed and understocked stands. The other analysed site and stand parameters individually had no significant influence on mistletoe incidence. The correlation of silver fir mortality in 2004 and mistletoe incidence in 2002/03 was analysed, and their strong correlation was established. Mistletoe could be considered as a bioindicator of silver fir decline, and probably a significant contributor to that decline. In the areas where mistletoe incidence is great it can be presumed that silver fir is significantly damaged.     

Structural changes in the vascular bundles of light-exposed and shaded spruce needles suffering from Mg deficiency and ozone pollution

The correlation between structural changes of the vascular bundles and needle yellowing was examined for needles of damaged spruce (Picea abies (L.) Karst.) growing at a Mg-deficient and ozone polluted mountain site in the Central Black Forest (840m a.s.l.). In the previous year's sun-exposed needles, the following sequence of events was observed: (1) rapid needle yellowing, (2) hypertrophy and anomalous divisions of cambium cells, (3) phloem collapse, and, (4) production of atypical xylem tracheids. Under defined shade (reduction of the photosynthetically active photon flux density of the ambient light by 85-90%), the needles remained green, while the phloem collapsed completely within the first 6 weeks of shading; subsequently, a reversal of the collapse was observed. Under both light conditions, the content of Mg not bound to chlorophyll (Mg(free)) was in the range of 0.1 mg g(-1) needle dry matter, and hardly changed throughout the investigation period. After Mg fertilization, the Mg(free) level of the previous year's needles increased to 0.2 mg g(-1) dry matter, the light-exposed needles remained green, and the vascular bundles developed no anomalies. The data show that the rapid needle yellowing of ozone-exposed Mg-deficient needles did not depend on the collapse of the phloem. Mg deficiency played a key role in the development of anomalous vascular bundles under light, and also appears to explain the transient changes in sieve cell structure under shade. The role of Mg deficiency, rather than ozone pollution, in the damage of the sieve cells was confirmed in a long-term ozone exposure experiment with young clonal spruce growing under defined conditions.     

Aluminium and manganese and their relation to calcium in soil solution and needles in three Norway spruce (Picea abies,L. Karst.) stands of Upper Austria

Soil solution and needles of three mature spruce stands in Upper Austria were analysed in order to investigate the uptake and possible toxic effects of Mn and Al, as these two elements become highly mobilised in the soil due to increasing acidity. The Ca/Al molar ratio in the soil solution was below 0.2 in the most damaged stand during almost the whole vegetation period. Despite different dynamics, Al reaches almost identical values in all stands at the end of the vegetation period in both 1-year (current) and 2-year-old needles, respectively. Therefore, needle analysis is not a useful tool for estimation of free Al in the soil. Needle contents of other elements could provide a better information for understanding the forest decline. Mn in the needles correlates significantly with Mn concentrations in the soil solution. As soil Mn will be mobilised by acidic input, Mn needle content can increase to very high levels. Manganese distribution, its interaction with calcium, and possible toxic effects are discussed.     

The ion concentration of dew condensed on Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.) needles

Summary Dew droplets collected with pipettes from coniferous needles were analysed for their ionic composition. Almost all samples of dew taken from Scots pine trees (Pinus sylvestris) showed significantly higher ion concentrations than those taken from Norway spruce trees (Picea abies). This can be explained by the micromorphology of the needle surface. The higher microscale roughness of the wax layer of a pine needle causes a more efficient flux of atmospheric aerosol particles compared to the spruce needle surface. Dew on coniferous needles is shown to be capable of maintaining pH values below 3 for several hours.     

Activity and activation state of ribulose-1,5-bisphosphate carboxylase of spruce trees with varying degrees of damage relative to the occurrence of novel forest decline

The aim was to determine whether a reduced carboxylation efficiency in needles of damaged spruce trees (Picea abies), is derived from a direct impairment of the ribulose-1,5-bisphosphate carboxylase (RuBP carboxylase) or there is an indirect inhibition of the RuBP carboxylase. In 1985, 1986 and 1987 measurements of RuBP carboxylase activity were carried out at three locations. Trees of different ages and degrees of damage were examined. RuBP carboxylase was assayed using both a rapid extraction method to determine the initial activity and an in vitro test after total activation to determine the total activity. The activation state was calculated as the ratio of initial activity to total activity.Within three vegetation periods the total activity in needles of damaged and apparently healthy or slightly damaged spruce trees indicated no definite difference in the annual average. On the other hand, in damaged needles a continued decline of the actual activation of RuBP carboxylase was established. The observation of continued depression of the activation state of the enzyme in needles of damaged spruce trees can possibly be due to a reduced photosynthetic electron transport rate.The measurements of the soluble protein content indicate a tendency to increased amounts in the needles of damaged trees. In accordance, a considerable increase of the activity of some enzymes like glutamine synthethase, phosphoenol-pyruvate carboxylase, and catalase could be noticed. However, there is no clear connection between the RuBP carboxylase and the content of soluble proteins.Abbreviations chl chlorophyll a+b, dw-dry weight, i.a-initial activity - P-700 reaction center of photosystem I - PVP polyvinylpyrrolidone 25 - RuBP ribulose-1,5-bisphosphate - RuBPCase ribulose-1,5-bisphosphate carboxylase - t.a. total activity     

Investigations on the Starch Content and Ultrastructure of Spruce Needles Relative to the Occurrence of Novel Forest Decline

The starch content and ultrastructure of needles of Norway spruce trees [Picea abies (L.) Karst.] taken from three natural habitats, undamaged or with symptoms of novel forest decline, were investigated during the course of three years. The starch content was clearly dependent on the seasons, with a maximum in spring and a decline during summer and autumn, leading to a minimum in winter. Needles of damaged trees from one habitat exhibited in all three years from August to October a significantly higher starch content than their undamaged counterparts. Microscopic investigations of these needle samples exhibited severe damage symptoms to the phloem in macroscopically green needles, though more intensely in the needles with yellowing symptoms. The phenomenon of starch accumulation is interpreted as a delay in starch mobilization, caused by a reduction in assimilate transport capacity of the needle phloem. The lower degree of starch accumulation in needles of damaged trees from a second natural habitat corresponded well to the lesser extent of phloem damage. The difference in damage patterns are discussed in terms of the differing air pollution situations acting on both habitats.