Cytology of Norway spruce needles

The structural changes in the albuminous, endodermal and mesophyll cells in the ageing needles of different aged healthy spruces (Picea abies [L.] Karst.) were studied on material collected in Germany and in Finland in winter. The results are considered as background information for the studies dealing with the structural changes in spruces suffering from biotic or abiotic stresses e. g., from the polluted air. Both light and electron microscopes were used.     

Fungi on spruce needles in Norway II

Tiarosporella parca has been found in different plots in Norway, mostly in small amounts. However, in Pasvik, East-Finnmark, this species was rather common and replaced the otherwise most common species, Lophodermiumpiceae.     

Life cycle of Lophodermium piceae in Norway spruce needles

The spruce needle inhabiting ascomycete Lophodermium piceae can live symptomless for several years, before it completes its life cycle within less than one year as detailed studies have shown.     

Effect of mineral nutrition content on oxygen exchange and chlorophyll a fluorescence in needles of Norway spruce

Photosynthetic O(2) evolution at high irradiances (approximately 600-1000 micro mol m(-2) s(-1)) and O(2) uptake in darkness were measured in needles of control, irrigated and irrigated-fertilized trees of Norway spruce (Picea abies (L.) Karst.). Measurements were made at 20 degrees C and at high CO(2) concentrations. The results suggest that, at given times of the year, a major part of the variation in gross photosynthesis of current-year and one-year-old needles across treatments is associated with differences in needle N content. Furthermore, the rate of O(2) uptake measured after 5 or 10 min in darkness was positively correlated with both the preceding rate of gross O(2) evolution and the N content in fully expanded current-year needles. Measurements of chlorophyll a fluorescence, taken simultaneously with measurements of O(2) evolution in current-year sun needles, showed that Stern-Volmer quenching of minimum fluorescence and the ratio of variable to maximum fluorescence in the dark- and light-adapted state were strongly correlated with the gross rate of O(2) evolution. This suggests that the increased rate of gross photosynthesis in needles of irrigated-fertilized trees was associated with adjustments in the thermal energy dissipation within photosystem II.     

Effect of thinning on anatomical adaptations of Norway spruce needles

Conifers and other trees are constantly adapting to changes in light conditions, water/nutrient supply and temperatures by physiological and morphological modifications of their foliage. However, the relationship between physiological processes and anatomical characteristics of foliage has been little explored in trees. In this study we evaluated needle structure and function in Norway spruce families exposed to different light conditions and transpiration regimes. We compared needle characteristics of sun-exposed and shaded current-year needles in a control plot and a thinned plot with 50% reduction in stand density. Whole-tree transpiration rates remained similar across plots, but increased transpiration of lower branches after thinning implies that sun-exposed needles in the thinned plot were subjected to higher water stress than sun-exposed needles in the control plot. In general, morphological and anatomical needle parameters increased with increasing tree height and light intensity. Needle width, needle cross-section area, needle stele area and needle flatness (the ratio of needle thickness to needle width) differed most between the upper and lower canopy. The parameters that were most sensitive to the altered needle water status of the upper canopy after thinning were needle thickness, needle flatness and percentage of stele area in needle area. These results show that studies comparing needle structure or function between tree species should consider not only tree height and light gradients, but also needle water status. Unaccounted for differences in needle water status may have contributed to the variable relationship between needle structure and irradiance that has been observed among conifers.     

Histology of magnesium-deficient Norway spruce needles influenced by nitrogen source

Effects of magnesium deficiency and variation in nitrate to ammonium ratio on needle histology and chlorophyll concentration were investigated in current-year and one-year-old needles of clonal Norway spruce trees (Picea abies (L.) Karst.). Six-year-old trees were grown for one year in sand culture with circulating nutrient solutions containing a sufficient (0.2 mM) or a limiting (0.04 mM) concentration of Mg. The nitrogen concentration was not varied (5 mM), but the NO(3) (-)/NH(4) (+)-ratio was adjusted to 0.76 in Mg-sufficient and to 1.86, 0.76 or 0.035 in Mg-limited plants. Visible symptoms of Mg deficiency occurred only in current-year needles, indicating adequate Mg nutrition before the experiment. Under conditions of Mg limitation, chlorophyll and Mg concentrations were lowest in needles of trees supplied with NH(4) (+) as the major nitrogen source and highest in needles of trees supplied with NO(3) (-) as the major nitrogen source. In current-year and one-year-old needles, starch accumulation induced by Mg deficiency was increased when NH(4) (+) was the major nitrogen source. The accumulation of tannin spherules in current-year needles, which occurred in response to Mg deficiency, also increased with decreasing NO(3) (-)/NH(4) (+)-ratios. Deficient Mg supply caused premature aging in tissues of the vascular bundle, as indicated by modifications of the cambium and increased amounts of collapsed sieve cells. The number of collapsed sieve cells was slightly lower in needles grown in a NH(4) (+)-dominated nutrient regime than in needles grown in a NO(3) (-)-dominated nutrient regime. We conclude that was not directly toxic to Norway spruce trees at the applied concentrations. However, effects of Mg deficiency were considerably greater in an NH(4) (+)-dominated nutrient regime than in a NO(3) (-)-dominated nutrient regime.     

Seasonal patterns of colonization of Norway spruce needles by Lophodermium piceae

Summary Colonization of living Norway spruce needles by Lophodermium piceae was studied in southern Sweden. One‐month to 3‐year‐old needles were assessed for frequency of infection. Needles appearing in the growing season of 1996 were monitored and sampled at 1 to 4 week intervals over a period of 14 months. Infection frequency of the current‐year needles increased from around 20% in July to approximately 40% in August 1996. This level had then doubled to approximately 80% after another growing season in August 1997. Multiple infections were rare in the needles during the first growing season, but after the second growing season at least 70% of the needles had multiple infections. The average number of infections in 9‐, 21‐ and 33‐month‐old needles were 1.3, 6.3 and 8.4, respectively. The size of the individual mycelia was estimated to be small in all needles, none of the individuals could be isolated from more than two adjacent needle sections, about 1.5 mm long. In conclusion, the number of L. piceae infections per needle increased during ageing of needles but mycelial growth is probably postponed until needle senescence and death.     

Structural features of water-soluble arabinogalactans from Norway spruce and Scots pine heartwood

 Isolated water-soluble acidic arabinogalactans from Norway spruce and Scots pine heartwood were analysed and compared to Siberian larch heartwood arabinogalactans. The carbohydrate monomer composition was determined by acid methanolysis and gas chromatography, while structural studies were performed by ¹³C NMR spectroscopy and methylation analysis. The main structural features were found to be the same in the three types of arabinogalactans. However, the structure of the arabinogalactans from spruce and pine were found slightly different from the structure of larch arabinogalactans. The amount of single unit side-chains, consisting of arabinose and glucuronic acid units, was higher in the spruce and pine arabinogalactans than in the larch arabinogalactans. The amount of glucuronic acid was higher in the spruce arabinogalactans than in the pine arabinogalactans. The pine arabinogalactans had a higher amount of side chains with more than two sugar units than the spruce arabinogalactans. Received 30 December 1999     

Effects of manganese deficiency on soluble apoplastic peroxidase activities and lignin content in needles of Norway spruce (Picea abies)

Apoplastic peroxidase activities were investigated in manganese-deficient and manganese-sufficient needles of field-grown Norway spruce trees (Picea abies L.). In Mn-sufficient needles, two sets of peroxidases, one with an alkaline pI >/= 9 and another with an acidic pI 相似文献   

Vascular tissue anatomy of Norway spruce needles and twigs in relation to magnesium deficiency

Vascular tissues of needles and twigs of Norway spruce with low foliar magnesium concentrations were examined by light microscopy. Observations were made on samples from (1) apparently healthy trees, (2) trees exhibiting progressive symptoms of decline, including the yellowing and fall of needles and the death of twigs and branches in the subapical canopy, and (3) trees recovering from symptoms of decline after fertilization with magnesium sulfate. Abnormalities in cambium and phloem anatomy, which were apparent in 2-year-old needles of declining trees, were seen only in needles at least 4 years old in healthy trees, suggesting that needles of declining trees are susceptible to precocious aging. Abnormalities in xylem anatomy were observed in needles 3 years or more in age in declining trees, but were never seen in needles of healthy trees. Over time, anatomical abnormalities in declining trees were seen in progressively younger twigs, which may explain the acropetal development of decline symptoms. After fertilization with magnesium sulfate, normal phloem and xylem were observed in the newly formed tissues.     

Relationships between environmental factors and height growth and yield of Norway spruce stands: a factor-analytic approach

Soil samples were taken from 148 experimental plots and subjectedto analysis of 34 morphological, physical and chemical properties.At the same time, eight climatic and topographic characteristicswere recorded. For each plot the values of top height and yieldwere available. The relationships between the soil propertiesand climatic/topographic characteristics were studied separately,for both sets of variables, by means of factor analysis. Theorthogonal varimax prerotation and the oblique promax rotationwere used. For soil characteristics, five common factors wereidentified, explaining 90 per cent of the total variance, relatedto soil acidity, water retention, humus content, nutrients andsoil texture. For climatic/topographic characteristics, threefactors were found, related to temperature, precipitation andterrain formation, explaining 85 per cent of variance. The relationships between the identified factors and heightgrowth and/or yield were quantified by polynomial regressionanalysis. Appropriate regression models were found for all factorsexcept the factor ‘nutrients’. Most regressionsproved to be non-linear. The applicability of the factor analysisfor determining the environmental factors is discussed.     

Lophodermium piceae and Rhizosphaera kalkhoffii in fallen needles of Norway spruce (Picea abies)

Falling needles collected from under individual 70-year-old Norway spruce (Picea abies) trees for about four years were checked for fungal fruitbodies. The most common fungi were Lophodermium piceae and Rhizosphaera kalkhoffii. Maximum frequencies of fruitbodies occurred in late autumn for L. piceae (ca. 80% of the needles), in summer for R. kalkhoffii (ca. 60%) and in winter for the third most common fungus, Tiarosporella parca (ca. 10%). The frequencies of needles with fruiting fungi varied greatly within and between years. This is the first report of T. parca from Sweden.     

A method for quantifying changes in the epicuticular wax structure of Norway spruce needles

A method for quantifying changes in the epicuticular wax structure of spruce needles (Pieea abies) has been developed for the scanning electron microscope. This evaluation system, based on a five-point scale, classifies only the epistomatal waxes of current-year needles. Two criteria, the different crystal wax forms and the amount of changed wax as a percentage of the total stomatal area have been used for classification. Statistically significant results (accuracy below ± 10% at 5% error level) were obtained under the following conditions. Mixed needle samples were taken from two adjacent trees per site. These were in a dominant and predominant social position in the stand. The needles were collected from the 7^th whorl of sun crowns exposed to all wind directions. The wax quality was determined by evaluating three stomata per needle. One stoma was investigated at the needle base, one m the middle and the third at the apex of the needle. 24 needles were evaluated per site. SEM-Investigations of needle waxes of 25 Austrian forest sites with no local emission source (background sites) showed, that the mean wax quality ranged from 1.6 to 4.1, according to this classification method. Stepwise multiple regression analyses with site characteristics and element contents of the needles (nutrient elements, heavy metals) confirmed correlations between the wax quality and the manganese- and sulphur-content of the needles and also the average degree of crown defoliation.     

Effects of simulated acid deposition on the surface structure of Norway spruce and silver fir needles

Treatments with acidified solutions of pH 3.5 cause alterations in the wax structure of the needles of silver fir and Norway spruce. The wax structures, especially those of the stomata, once they have been damaged, are not capable of complete reorganization even if the needle is repeatedly treated with distilled water.     

Characterization of glutathione S-transferases in needles of Norway spruce trees from a forest decline stand

Glutathione S-transferases (GST) detoxify many electrophilic xenobiotics, including several volatile organic compounds and pesticides. The GST activity for the conjugation of several xenobiotic substances was isolated from needles of Norway spruce (Picea abies L. Karst.) trees from a forest decline stand in the northern alps. Trees that exhibited different degrees of damage were selected from several stands in an altitude profile. The GST activity toward 1-chloro-2,4-dinitrobenzene (CDNB) in crude protein extracts of needles showed a seasonal pattern with highest activity during summer. The GST activity exhibited a strong dependence on the altitude of the stand showing highest activities in trees growing in the valley and lowest activities in trees growing in the summit regions of the mountain. When cytosolic GST from needles of healthy and damaged trees was purified, trees of healthy appearance exhibited three distinct GST isozymes with activities for the conjugation of CDNB and 1,2-dichloro-4-nitrobenzene (DCNB), whereas severely defoliated trees exhibited four GSTs with additional activity for the conjugation of ethacrynic acid. The main GST isozymes catalyzing the conjugation of CDNB differed in molecular weight, isoelectric point and catalytic properties between damaged and healthy trees.     

Interactions between soil scarification and Norway spruce seedling types

Interactions between scarification treatments and seedling types of Norway spruce were examined at two different locations in southern Sweden. The scarification treatments were not scarified control, mounding and soil inversion and the seedling types were a 10 week-old containerized seedling (mini seedling), a 2 year-old containerized seedling and a 2 year-old hybrid seedling, grown as a containerized seedling the first year and as a bare-root seedling the second year. Site preparation intensity had a greater effect on the mini seedlings. The results indicated that mini seedlings established faster in the soil inversion treatment compared to the larger seedling types. At the same seedling age, the mini seedlings had faster or similar growth rate as the containerized and the hybrid seedlings regardless of scarification treatment. Although mini seedlings grew as well as or even better than larger seedlings if they were successfully established, they were more sensitive to their planting environment and proper handling was critical. Problems with frost heaving and competing vegetation have to be taken into consideration when choosing site preparation method and seedling type.     

Tree- and needle-age-dependent variations in antioxidants and photoprotective pigments in Norway spruce needles at the alpine timberline

To cope with environmental stress, plants are equipped with antioxidative (e.g., ascorbate, glutathione and alpha-tocopherol) and photoprotective (e.g., xanthophyll cycle pigments) defense systems. We investigated the defense capacities of three tree age classes (mature, sapling and seedling) of Norway spruce (Picea abies (L.) Karst.) at a field site near the timberline. Biochemical data were expressed on both a needle dry mass and a surface area basis. Compared with current-year needles, previous-year needles contained higher mass- and area-based concentrations of chlorophylls and alpha-tocopherol, and a larger xanthophyll cycle pool that was in a more epoxidized state. Total glutathione concentration was lower, the glutathione pool was more reduced and the ascorbate pool was more oxidized in previous-year needles than in current-year needles. Needle concentrations of glutathione and alpha-tocopherol increased and chlorophyll concentration decreased with increasing tree age when expressed on a surface area basis. On a dry mass basis, these trends were reversed or nonexistent. The ascorbate pool was more reduced and the glutathione pool was more oxidized in needles of mature trees than in needles of saplings and seedlings. The proportion of protective xanthophyll cycle pigments decreased and the de-epoxidation state increased with increasing tree age. We conclude that tree age and the basis of expression of antioxidant concentration--surface area or dry mass--are important in scaling from seedlings to large trees.     

The influence of magnesium deficiency on carbohydrate concentrations in Norway spruce (Picea abies) needles

Both short- and long-term effects of Mg deficiency on carbohydrate metabolism were investigated in 6-year-old clonal Norway spruce (Picea abies (L.) Karst.) trees cultivated in sand culture with an optimal supply of nutrients, except for Mg which was supplied at 0.203, 0.041 and 0.005 mM to provide optimal, moderately deficient and severely deficient Mg supply, respectively. Annual changes in carbohydrate concentrations (starch, sucrose, glucose and fructose) were analyzed and diurnal changes were investigated on a single day during the summer. Older needles of trees in the moderate Mg-deficiency treatment developed tip-yellowing symptoms, whereas current-year needles remained green. The severe Mg-deficiency treatment led to pronounced yellowing symptoms in needles of all ages. Increased carbohydrate concentrations were observed before needle yellowing occurred. Diurnal and annual changes in carbohydrates were similar in all treatments; however, carbohydrate concentrations were influenced by Mg supply. In both Mg-deficiency treatments, starch concentrations increased in needles, especially during summer and autumn. Starch accumulation was more pronounced at the beginning of the Mg-deficiency treatments than at the end of the treatments. Sucrose, and to a minor extent, glucose and fructose concentrations tended to increase in response to Mg deficiency. The consequences of Mg deficiency on carbohydrate metabolism are discussed with respect to reduced plant growth and decreased transport rates of carbohydrates to sink organs.     

Co‐occurrence of the ascomycete Lophodermium piceae and the rust fungus Chrysomyxa abietis in Norway spruce needles

The frequency of needles and the proportion of needle segments infected by Lophodermium piceae were compared in symptomless and Chrysomyxa abietis‐infected, 1‐year‐old needles of Picea abies. In late spring, symptomless needles from both rust‐infected and healthy saplings were sampled. In addition, rust‐infected, totally chlorotic needles and needles with chlorosis along about half their length from the diseased trees were examined. In all three stands, the proportion of segments infected by L. piceae was larger in the rust‐infected half of the needle than in the symptomless half; but the difference was statistically significant in only one of the stands. The proportion of L. piceae‐infected segments among the nonrust‐infected needles was the same as that found for the uninfected half of rust‐infected needles (after correction for size differences). No differences in the proportion of L. piceae‐infected segments were found between the totally chlorotic, rust‐infected needles and the green needles of diseased or healthy trees.     

Relationships between crown projected area and components of above-ground biomass in Norway spruce trees in even-aged stands: Empirical results and their interpretation

Sample tree material was reanalyzed in order to study the relationships between horizontal crown projected area and components of above-ground biomass in Norway spruce (Picea abies (L.) Karst.) trees growing in even-aged stands. The needle mass of dominant trees increased linearly with the increase in crown projected area, but in co-dominant and dominanted trees the increase in needle mass levelled off toward larger crown projected areas. The branch mass of dominant and co-dominant trees accumulated faster than linearly with increasing crown projected area, whereas in dominated trees an approximately linear relationship existed between these two variables. The increase in needle and branch mass per unit increase in crown projected area was highest in dominant trees and decreased to co-dominant and dominated trees, i.e. with tree position in the canopy. The stem mass accumulated obviously faster than linearly and similarly in all tree classes with the increase in crown projected area. The narrow crown shape indicated a high density of all components of above-ground biomass per unit of crown projected area.