The spermatogenous body cell of the coniferPicea abies (Norway spruce) contains actin microfilaments

Summary In conifer pollen, the generative cell divides into a sterile stalk cell and a body cell, which subsequently divides to produce two sperm. InPicea abies (Norway spruce, Pinaceae) this spermatogenous body cell contains actin microfilaments. Microfilament bundles follow the spherical contour of the body cell within the cell cortex, and also traverse the cytoplasm and enmesh amyloplasts and other organelles. In addition, microfilaments are associated with the surface of the body cell nucleus. The sterile stalk cell also contains microfilament bundles in the cytoplasm, around organelles, and along the nuclear surface. Within the pollen grain, microfilament bundles traverse the vegetative-cell cytoplasm and are enriched in a webbed cage which surrounds the body cell. Microfilaments were identified with rhodamine-phalloidin and with indirect immunofluo-rescence using a monoclonal antibody to actin. The majority of evidence in literature suggests that the spermatogenous generative cell in angiosperms does not contain actin microfilaments, so the presence of microfilaments within the spermatogenous body cell inP. abies appears to be a fundamental difference in sexual reproduction between conifers and angiosperms.     

Microtubules and microfilaments are both responsible for pollen tube elongation in the coniferPicea abies (Norway spruce)

Summary InPicea abies (Norway spruce), microtubules and actin microfllaments both form a dense matrix throughout the tube mainly parallel to the direction of elongation. In these conifer pollen tubes the organization of this matrix is different from that in angiosperms. This study tests our hypothesis that differences in cytoskeletal organization are responsible for differences in tube growth and physiology. Pollen grains were germinated in media containing cytoskeletal disrupters and analyzed for germination, tube length, tube branching, and tip swelling. Disruption of microtubules significantly inhibits tube elongation and induces tube branching and tip swelling. Tip swelling is probably caused by disruption of the microtubules in the tip that are perpendicular to the direction of elongation. Confocal microscopy indicates that colchicine and propyzamide cause fragmentation of microtubules throughout the tube. Oryzalin and amiprophosmethyl cause a complete loss of microtubules from the tip back toward the tube midpoint but leave microtubules intact from the midpoint back to the grain. Disruption of microfilaments by cytochalasins B and D and inhibition of myosin by N-ethylmaleimide or 2,3-butanedione monoxime stops tube growth and inhibits germination. Microfilament disruption induces short branches in tubes, probably originating from defective microfilament organization behind the tip. In addition, confocal microscopy coupled with microinjection of fluorescein-labeled phalloidin into actively growing pollen tubes indicates that microfllament bundles extend into the plastid-free zone at the tip but are specifically excluded from the growing tip. We conclude that microtubules and microfilaments coordinate to drive tip extension in conifer pollen tubes in a model that differs from angiosperms.     

Effects of ozone on starch accumulation in Norway spruce (Picea abies)

Summary Fumigation with 100 g/m³ and 200 g/m³ ozone in closed-top fumigation chambers induced starch accumulation in chloroplasts of Norway spruce. This accumulation was probably due to a partial inhibition of the starch translocation at night. The intensity of the effect was dependent on the season and the age of the needles. The accumulation was reversed in winter. It is therefore unlikely that such an effect has much significance for plant health.     

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.     

Plantlet regeneration via somatic embryogenesis from subcultured callus of mature embryos ofPicea abies (Norway spruce)

Summary Embryogenic callus was initiated from radicles of mature embryos removed from imbibed seeds (24 h). Embryogenic and other nonembryogenic types of callus proliferated on a modified half-strength Murashige-Skoog medium (MS) basal medium (BM) supplemented withmyo-inositol, casein hydrolysate (CH), L-glutamine (gln) and growth regulators kinetin (KN), N⁶-benzyladenine (BAP) each (20×10⁻⁶ M), 2,4-dichlorophenoxyacetic acid (2,4-D) (50×10⁻⁶ M) Embryogenic callus bearing suspensor-like cells in a mucilaginous gel matrix was isolated and maintained by subculture every 10 to 12 days on BM with KN, BAP each (2×10⁻⁶ M) and 2,4-D (5×10⁻⁶ M). Somatic embryos developed spontaneously from the callus on this medium at 23±1° C. Closer examination revealed that numerous polyembryonic clusters, comprised of elongated cells (suspensors) and small dense cells with large nuclei (somatic embryos), occurred in the viscous gel. When this enriched embryonal-suspensor mass was subcultured to low 2,4-D (1×10⁻⁶ M), globular embryos developed by 40 to 60 days. Upon transfer to a liquid medium without growth regulators, the embryos elongated and developed cotyledons and shoots with needles. Plantlet development was completed by 30 days in a basal medium without CH, gln and growth regulators. The total culture time was 150 days. Approximately 40±10 embryos were formed from 500 mg of initial callus. Somatic embryogenesis became aberrant if embryos remained attached to the callus mass and were not subcultured within 10 to 12 days according to the described protocol. Somatic embryos were encapsulated in an alginate gel and stored at 4° C for nearly two months without visible adverse effects on viability. Editor's Statement This paper presents advances in the in vitro regeneration of a commercially useful plant species from stored seeds. In addition, data is presented on short-term storage of the plantlets, and long-term proliferation of the embryonal mass in vitro.     

Multivariate interpretation of the foliar chemical composition of Norway spruce (Picea abies)

Twenty-four chemical elements were analysed by INAA, ICP-AES and CN in needles of Norway spruce (Picea abies (L.) Karst.). Branches were sampled from 54 trees on eight sites in Switzerland and South Germany. From each tree, twigs were sorted into the most recent four or five age classes and their needles analysed separately. All measured concentrations could be considered as log-normally distributed and statistical analyses were, therefore, performed on logarithms. Variance components were estimated by maximum likelihood and compared between elements. Non-essential elements varied more than essential nutrients (Mn was an exception). The sites and the age of the needles were the most important sources of variance. The interaction between site and age, the individual tree, the sampled branch and the residual variance were usually much smaller sources of variance. The effects of the most significant factors – site and age – were further described by principal components and cluster analyses. Mineral elements either increased or decreased with the age of the needles according to their mobility in the phloem. Two different components were identified in the effect of the sites: a geochemical component linked to soil pH and a climate component linked to altitude, temperature and precipitation. Multivariate statistics are discussed as a tool for the interpretation of complex interaction patterns between element concentrations in plants.     

Molecular characterization of a phosphoenolpyruvate carboxylase in the gymnosperm Picea abies (Norway spruce)

Phosphoenolpyruvate carboxylase (PEPC) genes and cDNA sequences have so far been isolated from a broad range of angiosperm but not from gymnosperm species. We constructed a cDNA library from seedlings of Norway spruce (Picea abies) and identified cDNAs coding for PEPC. A full-length PEPC cDNA was sequenced. It consists of 3522 nucleotides and has an open reading frame (ORF) that encodes a polypeptide (963 amino acids) with a molecular mass of 109 551. The deduced amino acid sequence revealed a higher similarity to the C3-form PEPC of angiosperm species (86–88%) than to the CAM and C4 forms (76–84%). The putative motif (Lys/Arg-X-X-Ser) for serine kinase, which is conserved in all angiosperm PEPCs analysed so far, is also present in this gymnosperm sequence. Southern blot analysis of spruce genomic DNA under low-stringency conditions using the PEPC cDNA as a hybridization probe showed a complex hybridization pattern, indicating the presence of additional PEPC-related sequences in the genome of the spruce. In contrast, the probe hybridized to only a few bands under high-stringency conditions. Whereas this PEPC gene is highly expressed in roots of seedlings, a low-level expression can be detected in cotyledons and adult needles. A molecular phyiogeny of plant PEPC including the spruce PEPC sequence revealed that the spruce PEPC sequence is clustered with monocot and dicot C3-form PEPCs including the only dicot C4 form characterized so far.     

Cloning, characterization and localization of three novel class III peroxidases in lignifying xylem of Norway spruce (Picea abies)

Plant class III peroxidases (POXs) take part in the formation of lignin and maturation of plant cell walls. However, only a few examples of such peroxidases from gymnosperm tree species with highly lignified xylem tracheids have been implicated so far. We report here cDNA cloning of three xylem-expressed class III peroxidase encoding genes from Norway spruce (Picea abies). The translated proteins, PX1, PX2 and PX3, contain the conserved amino acids required for heme-binding and peroxidase catalysis. They all begin with putative secretion signal propeptide sequences but diverge substantially at phylogenetic level, grouping to two subclusters when aligned with other class III plant peroxidases. In situ hybridization analysis on expression of the three POXs in Norway spruce seedlings showed that mRNA coding for PX1 and PX2 accumulated in the cytoplasm of young, developing tracheids within the current growth ring where lignification is occurring. Function of the putative N-terminal secretion signal peptides for PX1, PX2 and PX3 was confirmed by constructing chimeric fusions with EGFP (enhanced green fluorescent protein) and expressing them in tobacco protoplasts. Full-length coding region of px1 was also heterologously expressed in Catharanthus roseus hairy root cultures. Thus, at least the spruce PX1 peroxidase is processed via the endoplasmic reticulum (ER) most likely for secretion to the cell wall. Thereby, PX1 displays correct spatiotemporal localization for participation in the maturation of the spruce tracheid secondary cell wall.     

ABA consumption in norway spruce (Picea abies) and white spruce (Picea glauca) somatic embryo cultures

Summary We investigated abscisic acid (ABA) metabolism among Norway and white spruce somatic embryo cultures which exhibited differences in maturation response when placed on racemic abscisic acid [(±)-ABA]. Differences in metabolic rate among the spruce genotypes could affect the ABA pool available for the maturation process, and might therefore be responsible for the differences in maturation response. The production of cotyledonary (stage 3) somatic embryos in cultures (genotypes) of Norway spruce (PA86:26A and PA88:25B) and of white spruce (WS1F cryoD and WS46) was compared. In each species pair one of the two genotypes failed to show stage 3 embryo development (respectively, PA88:25B and WS46). The investigation of ABA metabolism of each species pair showed that no substantial differences in ABA consumption or in the production of metabolites occurred. In each case ABA was metabolized to phaseic acid and dihydrophaseic acid over the 42-day culture period, metabolites were recoverable from the agar-solidified medium, and the sum of residual ABA and metabolites were equivalent to the ABA initially supplied. The results indicate that the process of ABA metabolism occurs essentially independently of somatic embryo maturation. NRCC no. 37345.     

Effects of Cd and Zn on the development of annual xylem rings of young Norway spruce (Picea abies) plants

Three-year-old spruce (Picea abies) saplings were planted and cultivated for 2 years in pots with 3 1 substrate, consisting of a homogenized mixture of sand, peat and forest soil with a high organic content (volume ratio 11.52). This substrate was amended with 10–180 mol Cd [kg soil dry weight (DW)]^–1, 50–7500 mol Zn (kg soil DW)^–1 (determined with 1 M ammonium acetate extracts) or combinations of both elements. Annual xylem growth rings in stems of plants treated with 50 mol Cd (kg soil DW)^–1 or 7500 mol Zn (kg soil DW)^–1 were significantly narrower than in control plants. Growth reductions were more pronounced in the second year of the experiment. The contents of Cd and Zn in stem wood and needles were positively correlated with the substrate concentrations. The Mg contents of the spruce needles were inversely correlated with soil concentrations of Cd and Zn. Root development was impeded at moderate concentrations of Cd (50 mol kg^–1) or Zn (1000 mol kg^–1) in the substrate. The adverse effects of potentially toxic trace elements, like Cd or Zn, on xylem growth of spruce plants are discussed with regard to possible growth reductions in forest trees under field conditions.     

Dendrochronology of Norway spruce (Picea abies (L.) Karst.) from two range centres in lowland Poland

In Europe, spruce grows in two main regions which meet in Poland, one to the north and east, the other to the south and west. The northeastern area ranges from the northern treeline extending from Norway to Siberia, to southern Sweden, north-eastern Poland and the southern Ural mountains. The southwestern spruce region reaches the mountainous areas of the Alps and the Balkans, the mountains and uplands of the Sudety Mountains, the Carpathians and neighbouring lowlands. Opinions about the distribution of Norway spruce have changed over the years, and its scarcity in the centre of Poland has been strongly debated. The favoured current theory is that Norway spruce once had a continuous distribution in Poland. It is assumed that the rare occurrence in the central Polish lowland is due to a combination of unfavourable soil conditions and previous management activity. The main aim of this work was to analyse climate–growth relationships of Norway spruce in eastern Poland and distinguish regions with similar increment patterns with regards to spruce range. Spruce growth in northern Polish sites is positively correlated with rainfall from May to July. Tree-ring widths in southern sites are more correlated with March temperature. Selected homogenous regions are the same as range types. Trees from the so-called “spruceless area” seem to have similar climate–growth relationships to trees from the southern region. This finding does not settle the question of the origin of the trees, but it does indicate that similar environmental conditions exist in these two areas and proves that the climate was not a limiting factor there.

Fungal injury to seed tissues of Norway spruce,Picea abies (L.) Karst.

Norway spruce bore an abundance of cones in Finland in 2000, but these cones were often fungal-infected. The seeds had structural injuries that were revealed when seed samples were examined using light (LM) and a field emission scanning electron microscope (FESEM). Two main types of spores were found either in the tissues inside the seed coat or on the sarcotesta, the outermost layer of seed coat. The spores of Chrysomyxa pirolata appeared particularly in the nucellar tissue, where the cell walls were disintegrated at the middle lamellae and cytoplasm was disrupted. Degenerated remnants of fungal structures resembling aecial peridium were found close to aeciospores. The tissue of the megagametophyte differed also from that of a normal mature seed. Conidia of Thysanophora penicillioides were often encountered on the sarcotesta where the ordinary wax cover was missing. Fungal injury occurred in the nucellar layers that shelter the embryo and megagametophyte from desiccation and oxidation. Destruction of these structures together with rapid opening of the seed coat advance deterioration of seeds during storage and may cause unexpected economic losses in forest plant production.     

Developmental disorders in buds and needles of mature Norway spruce, Picea abies (L.) Karst., in relation to needle boron concentrations

Dieback of the terminal shoot and consequently bushy growth induced by boron deficiency have been reported widely throughout the world in several tree species. Recently, similar growth damage was documented in half of the young spruce stands in eastern Finland. To clarify the role of B deficiency, the light microscopic structure of emerging buds and of developing and previous-year needles of mature Norway spruce (Picea abies L. Karst.) from damaged (D stand), partly damaged (PD stand) and healthy (H stand) stands were analysed. The samples, on which needle nutrient concentrations were also determined, were taken seven times between early spring (April) and early winter (November). Cell death characterized by precipitation of the cell content, possibly due to the release of tannins after membrane rupture, was seen in the apex of emerging buds, and this led to fatal damage in about half of the buds in the trees from the D stand, where the needle B concentration was well below the deficiency level of 4–5 mg kg⁻¹. Furthermore, an increase in living cells that accumulated tannins in the vacuoles, which is a common stress and/or defense reaction, was found in the primordial shoots of buds and in the differentiating needles in the PD and D stands. The increase in the areas of the central cylinder and of the xylem found in the needles indicate structural plasticity during needle differentiation to drought. The time frame for bud emergence from late May up to mid-September means that an adequate B supply is necessary throughout the summer in order to avoid fatal bud damage and thus bushy growth of the trees.     

Isolation of two cytokinin metabolites from the rhizosphere of Norway spruce seedlings (Picea abies L. Karst.)

Roots of young Norway spruce seedlings were incubated under hydroculture conditions in a synthetic nutrient medium containing either ³H-isopentenyladenosine, isopentenyladenosine or zeatin riboside. When feeding with ³H-isopentenyladenosine a new radiaolabelled metabolite was found in the feeding solution as well as in root extracts. Isopentenyladenosine and zeatin riboside were metabolised and for both compounds an unknown metabolite was detected in the feeding solution. The metabolites were purified by solid phase extraction, HPLC and partially characterised. A major characteristic of the metabolites is their reactivity in the presence of NH₄OH, which results in the formation of the cytokinin bases isopentenyladenine or zeatin, respectively. UV-spectra and the chemical characteristics indicate that the new metabolites are closely related. The GC-MS analysis revealed, that the metabolites are true derivatives of isopentenyladenine and zeatin. The biogenesis of the new metabolites is discussed with regard to plant microbial interactions.Abbreviations Ck(s) = cytokinin(s) - GC-MS = gas chromatography-mass spectrometry - iP = isopentenyladenine - [9R]iP = isopentenyladenosine - [9G]iP = isopentenyladenine-9-glucoside - [9R-MP]iP = isopentenyladenosine-5-monophosphate - Z = trans-zeatin - [9R]Z = trans-zeatin riboside     

Diastereomeric stilbene glucoside dimers from the bark of Norway spruce (Picea abies)

As part of a long-term study of the chemical defenses of Norway spruce (Picea abies) against herbivores and pathogens, a phytochemical survey of the phenolics in the bark was carried out. Eight stilbene glucoside dimers, designated as piceasides A-H (1a-4b), were isolated as four 1:1 mixtures of inseparable diastereomers. Their structures were determined by extensive spectroscopic means including 1D (1H and 13C) and 2D NMR (1H-1H COSY, HSQC, HMBC, ROESY) spectra, and were supported by enzymatic hydrolysis and computational analysis.     

Tissue-specific expression of Pa18, a putative lipid transfer protein gene,during embryo development in Norway spruce (Picea abies)

A full-length Picea abies cDNA clone Pa18, encoding a protein with the characteristics of plant lipid transfer proteins, has been isolated and characterized. The size of the deduced 173 amino acid (aa) long protein is around 18 kDa. The first 100–120 aa show similarity to angiosperm lipid transfer proteins in amino acid sequence as well as in predicted secondary structure. The Pa18 gene is constitutively expressed in embryogenic cultures of Picea abies representing different stages of development as well as in non-embryogenic callus and seedlings. The Pa18 gene product has an antimicrobial activity. In situ hybridization showed that the Pa18 gene is equally expressed in all embryonic cells of proliferating embryogenic cultures but during embryo maturation the expression of the gene in maturing and mature somatic as well as in mature zygotic embryos is stronger in the outer cell layer than in other tissues. Southern blot analysis at different stringencies was consistent with a single gene with one or two copies rather than a gene family. Twenty independent transgenic sublines over- and under-expressing the Pa18 gene under the Zea mays ubiquitin promoter were established. There was a high yield of mature somatic embryos with a smooth surface only in untransformed, control cultures. Irrespective of the expression level of Pa18, the somatic embryos started to mature when given a maturation treatment. However, in the transgenic sublines, the outer cells in the maturing embryos frequently became elongated and vacuolated instead of remaining small and uniform. One explanation for this was that the expression of Pa18 was not restricted to the outer cell layer in transformed sublines. Angiosperms and gymnosperms separated about 300 million years ago and the embryo genesis is different in the two groups. The outer cell layer (protoderm), the first tissue to differentiate, is less clearly delineated in gymnosperms. For normal embryo development in angiosperms, expression of the LTP gene must be restricted to the protodermal cells. In this work we show that the expression of the Pa18 gene must be restricted to the putative protodermal cells of the gymnosperm.     

The significance of structure for imbibition in seeds of the Norway spruce,Picea abies (L.) Karst.

Since the observations of those regularly handling Norway spruce [Picea abies (L.) Karst.] seeds with regard to their imbibition frequently disagree with earlier opinions that this process is markedly inhibited by the seed coat, we decided to examine the morphological factors influencing imbibition in seeds of different colour and different provenances. The seed coat, consisting of the sarcotesta, sclerotesta and endotesta, was found to have little influence on the passage of water, despite the presence of sclereids full of wax lamellae. No differences in seed coat structure were observed between provenances or colours of seeds. The cells of the endotesta were lignified in the area of the micropyle, however, and stood out lip-like on the outer surface of the micropyle after imbibition. An opening in the sclerotesta filled with parenchyma cells was also seen at the chalazal end of the seed. Neither of these openings, which were covered by accumulations of wax, served as the main route for the passage of water, though the micropyle opened up slightly after only 24 h incubation, when the lignified cells bordering it swelled differently from the rest of the endotesta. The progress of water into the seed soon discontinued, however, as the tip of the nucellar cap, covered with wax and crystals, effectively plugged the micropyle. This opening of the micropyle may be the reason why the IDS method does not always succeed in separating viable from non-viable spruce seeds sufficiently well by their density. Imbibition was mostly regulated by the lipophilic layers surrounding the endosperm, which are mainly of nucellar origin, and particularly the megaspore membranes, the outer and inner exine. Imbibition was further hampered by the impermeable nucellar cap, which covered about 3/4 of the length of the endosperm and had merged with the outer exine at its edges. Deposits of wax were observed both between the exines and between the endotesta and the nucellar layers at the edges of the nucellar cap. Waxes may serve as a defence against diseases at the sites of water penetration, while simultaneously increasing the significance of the nucellar endosperm covers as regulators of imbibition.     

The allelic distribution at an acid phosphatase locus in Norway spruce (Picea abies) along similar climatic gradients

Summary The allele frequency distribution at a polymorphic acid phosphatase locus (APH-B) was determined in natural populations of Norway spruce (Picea abies) from a latitudinal transect in Finland, an altitudinal transect in the Austrian Alps, and from different locations of the Swiss range. The three independent population groups, selected with respect to similar climatic gradients, were studied to detect the forces that cause the geographic variation at the APH-B locus.In almost all of the populations investigated, four alleles (APH-B₁ — B₄) could be identified at this enzyme locus, however, the alleles b₁ and B₂, as well as B₃ and B₄, show a great similarity according to their phenotypic appearance after electrophoresis as well as to their frequency distributions along the different transects. With the aid of some theoretical considerations and data comparisons, a selective equivalence of the alleles B₁ and B₂, as well as B₃ and B₄, could be ascertained, thus reducing the number of alleles that can respond differently to natural selection.After combining the frequencies of the selectively equivalent alleles, similar clinal variation patterns could be observed along the different geographical transects, where-by the frequency of the allele group APH-B₁ /B₂ markedly increases with latitudes in Finland and towards higher elevations in the Alps. Correspondingly, the allele group APH-B₃/B₄ predominates in the southern parts of Finland and in the lowlands and foothills of Austria and Switzerland. It is therefore concluded that natural selection causes the geographic variation pattern at the APH-B locus and that one or several temperature variables function as an at least predominant selective force. Possible relationships between this enzyme polymorphism and other tree characters and the physiological role of acid phosphatases in tree adaptation were discussed.     

The seasonal variation of 20 elements in 1st and 2nd year needles of Norway spruce,Picea abies (L.) Karst

Summary The concentration of 20 elements (including eight nutritional elements) in spruce needles was monitored by taking samples at nine different dates in a single year. On each date the youngest and the 1-year-old needles were collected at four different sites. The needles were washed with an organic solvent to remove surface contamination before analysis by instrumental neutron activation. Although the sites showed widely different concentrations for the same element, they could be treated as a uniform population using normalized concentration values. Taking the time dependence of the concentration and the ratio of the concentrations in new and old needles as criteria, the following three groups of elements could be distinguished: 9 group I —Ca, Sr, Ba, and Mn; group II — Al, Br, Co, Fe, Hg, La, Sc, Sb, and Zn; group III — K, Rb, Cs, P, and Cl. Mg and Na did not fit into any of these groups. Within group I and III elements the strongest variations occur during the growing season (late spring and early summer) and the weakest during the dormant season; the variation is greater in the younger needles. Group II elements show an approximately linear increase throughout the year in both the younger and older needles. The concentrations of elements of groups I and II increase with time, while those of group III decrease. Na is unique insofar as its concentration is influenced by foliar uptake of road de-icing salt. The concentration of elements belonging to the same group shows strong inter-element correlations. The grouping of elements deduced here shows similarities with the chemical nature of the elements and with their transportability within the plant.     

Patterns and mechanisms of transpiration in a large subalpine Norway spruce (Picea abies (L.) Karst.)

In situ water relations of a large subalpine Norway spruce (Picea abies) were analyzed by simultaneous measurements of sap flow at different crown positions. In the diurnal scale, transpiration varied greatly, both spatially and temporally. Over longer periods, however, different parts of the crown transpired in fairly constant proportions. The average estimated transpiration was about 3.5 times greater in the upper than in the lower half and decreased 1.6-fold from south to north. Water intercepted from rain, fog and dew buffered and significantly decreased the transpiration. The effect was strongest in those parts which were least coupled to the free atmosphere. The top of the crown seemed to experience a regular shortage of water shortly after starting transpiration, when it was forced to switch from internal reserves to sources in the soil. Further, lower branches then started transpiring, which may have led them to compete for the water. An enhanced nocturnal sap flow during warm and dry winds (Foehn) indicated that the tree also transpired at night. Shaded twigs had more capacity to intercept water externally than twigs in the sun. The significance of the crown structure for interaction with water in both liquid and vapour phases is discussed.