Elongation and circumnutation oscillations of hypocotyl of pine seedlings (Pinus silvestris L.)

The elongation of pine seedlings (Pinus silvestris L.) is associated with nutation movements. Trajectories of these growth oscillations were recorded by film technique in horizontal and vertical projection during a three day period of growth. On the basis of these data the parameters of elongation and nutation oscillations,i.e. rate, amplitudes and frequency of oscillations, were calculated and their changes during plant development compared. Oscillation trajectories are circular or elliptic spirals the amplitudes of which are increasing with the age of hypocotyl from 1.5 mm to 7.5 mm. The frequencies of nutations are decreasing during the growth from 0.5 to 0.2 rev. h^?1. On the other hand, the growth rate of hypocotyl increased from values near to 10^?3 mm h^?1 at the beginning of the experiment to 4×10^?1 mm h^?1 recorded at the end of the third day. The zone of nutation curvature was slightly transferred from the middle of the hypocotyl toward the apex and its location has not been identical with that of elongation. This indicates that the system controlling nutation oscillations need not be identical with that controlling direction of elongation. At a certain stage of development behaviour of the decapitated pine hypocotyl is analogical to that of the root without the centre of georeception. A possibility of analogy of the system controlling direction of hypocotyl growth and of the system proposed for geotropical control of root growth is discussed.     

Nutation rhythm of growing pine hypocotyl (Pinus silvestris L.) interferred with a phototropic stimulus

The interaction of nutation oscillation and phototropic curvature mechanisms of growing pine hypocotyl(Pinus silvestris L.) was investigated. Nutation parameters of growing hypocotyl are not affected by continuous lateral illumination. Frequency and nutation amplitudes undergo changes which are dependent on plant development and are nearly the same when plants are illuminated from one side or from above. Lateral illumination, however, induces a phase shift. This deviation from a normal nutation rhythm usually disappears after a period of time equal to three nutation revolutions,i.e. after 12 h. The positive phototropio response of nutating hypocotyl is dependent on the phase in which the hypocotyl is exposed to the lateral illumination:(a) nutation perihelium,(b) ahelium and(c) nutation interphase. These response-variants are compared with the expected gradients of plant hormones at the opposite sites of the hypocotyle. Analysis of location of the individual movement reactions(i.e. oscillations as a component of the geocontrol system and phototropic curvature) together with the determination of the zone of maximum elongation indicates the existence of different receptory centres of these two movement systems. In the case of the phototropic response reaction it is possible to consider the presence of different photo- and biochemical mechanisms at least for a part of this system.     

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.     

Promotion of hypocotyl elongation in loblolly pine (Pinus taeda L.) by indole-3-acetic acid

Elongation of excised loblolly pine ( Pinus taeda L.) hypocotyls was promoted by indole-3-acetic acid and the fungal metabolite, fusicoccin. Gibberellic acid, kinetin, zeatin, or zeatin-riboside were either without effect or promoted elongation only slightly. The most auxin-responsive tissue was just below the cotyledonary node, and elongation was confined to sections excised from the upper 2 cm of the hypocotyl. Indole-3-acetic acid induced elongation rates in the hypocotyl sections equal to those of intact hypocotyls when the sections were excised from young seedlings. Elongation rates decreased in intact hypocotyls before the excised tissues lost responsiveness to the auxin. Hypocotyl elongation in loblolly pine is discussed in relation to hypocotyl elongation in angiosperm species.     

The effect of lateral illumination on growth oscillations of pine seedling hypocotyls (Pinus silvestris L.)

Phototropic sensitivity of forest wood seedlings to lateral illumination was proposed as an early assay for distinquishing various ecotypes of woody plants of the same species. Statistical analysis showed that results were significantly influenced by heterogenity of experimental material caused by an interference of phototropic movements and natural oscillations of hypocotyl. Both movements of pine seedlings (Pinus silvestris L.) were registered by phase photography and their mechanism was analyzed. The apical part of growing hypocotyl illuminated from above oscillates in a space spiral with frequency 3.3 h at mean growth rate 0.66 mm h^?1. The mean size of spiral amplitudes is 2.9 mm. The oscillation rhythm is disturbed after the lateral illumination and a phase shift was observed as a result. A new oscillation rhythm with frequency 3.9 h and mean growth rate 0.69 mm h^?1 was stabilized after a period of time equal to one nutation turn. Oscillation amplitudes were increased to 4.3 mm. In addition to the radiation intensity the appearance of the phototropic response to light pulse is first of all effected by the actual position of the apical part of hypocotyl in the course of endogenous circumnutations. A uniform plant material for the early assay may be obtained by selection of seedlings which are at the beginning of lateral illumination in the same phase of nutation rhythm. Under such conditions the deviations of longitudinal axis of oscillating spirals characterize the actual phototropic curvature.     

Gradients and dynamics of inner bark and needle osmotic potentials in Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies L. Karst)

Preconditions of phloem transport in conifers are relatively unknown. We studied the variation of needle and inner bark axial osmotic gradients and xylem water potential in Scots pine and Norway spruce by measuring needle and inner bark osmolality in saplings and mature trees over several periods within a growing season. The needle and inner bark osmolality was strongly related to xylem water potential in all studied trees. Sugar concentrations were measured in Scots pine, and they had similar dynamics to inner bark osmolality. The sucrose quantity remained fairly constant over time and position, whereas the other sugars exhibited a larger change with time and position. A small osmotic gradient existed from branch to stem base under pre‐dawn conditions, and the osmotic gradient between upper stem and stem base was close to zero. The turgor in branches was significantly driven by xylem water potential, and the turgor loss point in branches was relatively close to daily minimum needle water potentials typically reported for Scots pine. Our results imply that xylem water potential considerably impacts the turgor pressure gradient driving phloem transport and that gravitation has a relatively large role in phloem transport in the stems of mature Scots pine trees.     

Variation in fine root biomass of three European tree species: Beech (Fagus sylvatica L.), Norway spruce (Picea abies L. Karst.), and Scots pine (Pinus sylvestris L.)

Abstract

Fine roots (<2 mm) are very dynamic and play a key role in forest ecosystem carbon and nutrient cycling and accumulation. We reviewed root biomass data of three main European tree species European beech, (Fagus sylvatica L.), Norway spruce (Picea abies L. Karst.) and Scots pine (Pinus sylvestris L.), in order to identify the differences between species, and within and between vegetation zones, and to show the relationships between root biomass and the climatic, site and stand factors. The collected literature consisted of data from 36 beech, 71 spruce and 43 pine stands. The mean fine root biomass of beech was 389 g m^?2, and that of spruce and pine 297 g m^?2 and 277 g m^?2, respectively. Data from pine stands supported the hypothesis that root biomass is higher in the temperate than in the boreal zone. The results indicated that the root biomass of deciduous trees is higher than that of conifers. The correlations between root biomass and site fertility characteristics seemed to be species specific. There was no correlation between soil acidity and root biomass. Beech fine root biomass decreased with stand age whereas pine root biomass increased with stand age. Fine root biomass at tree level correlated better than stand level root biomass with stand characteristics. The results showed that there exists a strong relationship between the fine root biomass and the above-ground biomass.     

Distribution of eriophyoid mites (Acari: Eriophyoidea) on coniferous trees

The aim of the paper was to determine the infestation parameters and species composition of eriophyoid mites for different parts of Norway spruce and Scots pine as well as for different age groups of the trees. The observations on the occurrence of the mites were conducted in 2004 and 2005 in 4 locations distributed in various regions of Poland, accounting for 11 environments (location x year). Three plant age groups were studied: (1) adult trees: 40–60 years old, additionally divided into three levels: top, middle and bottom; (2) young trees: 6–10 years old; and (3) seedlings: 2–3 years old. The same number of species (five) occurred on each coniferous tree, but only one, the rarest, was common on both tree species. Out of 500 samples for each species, mites were found on 279 pine (55.8%) and 252 spruce samples (50.2%). No tendency for the mites to choose any particular level on Scots pine and Norway spruce was observed. In addition, no tendency for the mites to choose trees from any of the age groups was observed for both Scots pine and Norway spruce, in the latter case the result obtained also for mite species subdivided into vagrant and refuge-seeking ones. Final conclusions were that in case of adult trees samples can be taken from the bottom part of a tree; however, sampling from young trees growing among adult trees may be seen as the most efficient sampling method.     

Genetic information from progeny trials: a comparison between progenies generated by open pollination and by controlled crosses

When conifer progenies generated by open pollination are assessed in field tests, it is usually assumed that all progenies of the same mother are true half-sibs. This assumption may be invalid, leading to overestimation of additive genetic variation and heritability and to biased breeding values. From one Scots pine (Pinus sylvestris) and one Norway spruce (Picea abies) seed orchard, containing 28 and 36 parent clones, respectively, progenies generated by open pollination (OP) and by controlled crosses (CC) were planted in adjacent trials at two to three sites in southern Sweden. The tree height and diameter at breast height were measured, and genetic parameters based on these traits were estimated for OP and CC progenies separately, in order to enable comparisons. Narrow-sense heritability estimates for Scots pine and Norway spruce OP progenies (in the ranges 0.04–0.13 and 0.15–0.38, respectively) did not differ significantly from CC estimates (0.07–0.12 and 0.23–0.30), suggesting that OP-based heritability values were not overestimated to any great extent. Similarly, genetic correlations between OP and CC progenies were in the ranges of 0.87–0.88 and 0.74–0.77 for Scots pine and Norway spruce, respectively, being significantly lower than unity only in the case of Norway spruce. OP-based breeding values for both species should therefore correspond well with those predicted from CC progenies, albeit not perfectly for Norway spruce. In conclusion, the assumption of true half-sibs for OP progenies was not violated to the extent that genetic parameter estimates or breeding value predictions were seriously biased.     

Differential response of Scots pine seedlings to variable intensity and ratio of red and far‐red light

We investigated the response to increasing intensity of red (R) and far‐R (FR) light and to a decrease in R:FR ratio in Pinus sylvestris L. (Scots pine) seedling. The results showed that FR high‐irradiance response for hypocotyl elongation may be present in Scots pine and that this response is enhanced by increasing light intensity. However, both hypocotyl inhibition and pigment accumulation were more strongly affected by the R light compared with FR light. This is in contrast to previous reports in Arabidopsis thaliana (L.) Heynh. In the angiosperm, A. thaliana R light shows an overall milder effect on inhibition of hypocotyl elongation and on pigment biosynthesis compared with FR suggesting conifers and angiosperms respond very differently to the different light regimes. Scots pine shade avoidance syndrome with longer hypocotyls, shorter cotyledons and lower chlorophyll content in response to shade conditions resembles the response observed in A. thaliana. However, anthocyanin accumulation increased with shade in Scots pine, which again differs from what is known in angiosperms. Overall, the response of seedling development and physiology to R and FR light in Scots pine indicates that the regulatory mechanism for light response may differ between gymnosperms and angiosperms.     

Levels of damage of Scots pine and Norway spruce caused by needle miners along a SO2 gradient

Needle damages caused by mining insects on Scots pine and Norway spruce were studied in the vicinity or a pulp mill. The abundance of needles mined by the pine bud moth Exoteleia dodecella (Lepidoptera: Gelechiidae) on Scots pine Pinus sylvestris , and the spruce needle miner Epinotia tedella (Lepidoptera: Tortricidae) on Norway spruce Picea abies had a significantly peaked response curve on logarithmic distance scale. For pine bud moth, the maximum expected population density was estimated to be at the distance of 1.35 km from the factory. The maximum expected population density for the spruce needle miner was at the distance of 1.53 km from the factory. However, for both species the curves were significantly different among transects. Both species had a peaked and significant response to sulphur level in needles as well. The maximum expected density in pine was at 1270 ppm, and in spruce at 1070 ppm sulphur concentration in pine needles. The results are consistent with earlier reports demonstrating that these mining insects frequently attack trees suffering from air pollution. The nonlinear response of both species to distance from the pulp mill suggests that E. dodecella on pine and E. tedella on spruce are rather indicators of the zone of intermediate air pollution than of strongly polluted or nearly unpolluted sites. This also agrees with the plant stress-insect performance hypothesis indicating that insect response varies with the magnitude of stress, and at very high stress levels a tree no longer provides the insects with relevant food.     

Selective stripping of rowan (Sorbus aucuparia L.) bark by cattle in North-east Scotland

Summary

During winter 1973–4 beef cattle removed bark selectively and extensively from rowan growing in two Aberdeenshire woods. Some bark was also taken from Norway spruce and willow, but damage to all other species, including ash, beech, birch, gean, larch, Scots pine and sycamore, was negligible. In other woodlands utilised by cattle in this area bark-stripping was either less severe or absent. Possible causes for the bark-stripping are discussed. The rowan bark was not distinctive in cover, species of epiphytic lichen or inorganic composition, except perhaps for magnesium.     

Purification of Two Superoxide Dismutase Isozymes and Their Subcellular Localization in Needles and Roots of Norway Spruce (Picea abies L.) Trees

Two isozymes of superoxide dismutase (SOD; EC 1.15.1.1) were purified from Norway spruce (Picea abies L.) needles to apparent electrophoretic homogeneity. Purification factors were 354 for SOD I and 265 for SOD II. The native molecular mass of both purified enzymes was approximately 33 kD, as determined by gel filtration. The subunit molecular weights, as estimated by sodium dodecyl sulfate polyacrylamide gel electrophoresis, were 20,000 for SOD I and 16,000 for SOD II in the presence of 2-mercaptoethanol, and 15,800 and 15,000, respectively, in its absence. These results indicate that the native enzymes were homodimers whose subunits contained intrachain disulfide bonds. Isoelectric points determined by nondenaturing isoelectric focusing were 4.5 and 5.5 for SOD I and II, respectively. NH₂-terminal sequence analysis of the first 22 to 23 amino acids revealed 70 to 75% sequence identity with chloroplastic CuZn SODs from other plant species for SOD I, and 75% sequence identity with the cytosolic CuZn SOD from Scots pine for SOD II. SOD I was the major activity in needles and it was associated with chloroplasts. SOD II activity was dominant in roots.     

Single-cell origin and development of somatic embryos in Picea abies (L.) Karst. (Norway spruce) and P. glauca (Moench) Voss (white spruce)

The origin and development of somatic embryos in calli initiated from immature zygotic embryos of Picea abies (L.) Karst. (Norway spruce) and P. glauca (Moench) Voss (white spruce) was studied. Immature zygotic embryos cultured on callus induction medium produced two types of white calli that were phenotypically different from one another. The callus that proliferated from the hypocotyl region was white to translucent, glossy, mucilaginous and embryogenic. The callus mass which originated from the radicle end was reddish-white, nonmucilaginous and nonembryogenic. Whole mount preparations of the entire explant with two different types of calli showed the presence of embryogenic cells in the mucilaginous callus mass derived from the hypocotyl region of the zygotic embryo. The origin of somatic embryos in both Norway and white spruce could be traced to single cells of the hypocotyl callus.Abbreviations 2,4-D 2,4-dichlorophenoxyacetic acid - BAP 6-benzylaminopurine     

Patterns of genetic parameters for height in field genetic tests of Picea abies and Pinus sylvestris in Sweden

We reviewed the genetic parameter estimates carried out from 1992 to 2006 for height increment in genetic tests of Norway spruce and Scots pine, to describe patterns of genetic variation, heritability, and genetic correlations. The material included seedling and clonal tests in Sweden, aged between 5 and 20 years. Multiple regression was used to explore relationships between parameter values and test environments. Results showed moderate narrow-sense heritabilities ([^(h)]² {\hat{h}^2} : mean =0.29 in Norway spruce; mean =0.23 in Scots pine) that decreased with test site latitude for both species. In Norway spruce, [^(h)]² {\hat{h}^2} increased with better growth and decreased with tree age, while for Scots pine, [^(h)]² {\hat{h}^2} increased with tree age and southward transfer. The additive genetic coefficient of variation (; mean 15%), in Norway spruce, decreased with growth as well as site latitude. in Scots pine (mean =8.5%) increased with southward transfer and more southerly test latitude. Additive and genotypic within-site genetic age-age correlations in Norway spruce were high, with mean r _A and r _G of 0.92 and 0.85, respectively. Corresponding across-sites estimates were on average lower. Genetic parameters were better expressed on favorable sites, at younger ages in Norway spruce and at older ages in Scots pine. The results imply that gain calculations should be based on different parameters in the two species. For maximizing genetic gain in the Swedish breeding program, testing times could be shorter for Norway spruce than for Scots pine. The investigation showed a large variation in parameter estimates from different field experiments, highlighting the importance of testing over multiple sites.     

Sensitivity of French temperate coniferous forests to climate variability and extreme events (Abies alba,Picea abies and Pinus sylvestris)

Questions: (1) How do extreme climatic events and climate variability influence radial growth of conifers (silver fir, Norway spruce, Scots pine)? (2) How do elevation and soil water capacity (SWC) modulate sensitivity to climate? Location: The sampled conifer stands are in France, in western lowland and mountain forests, at elevations from 400 to 1700 m, and an SWC from 50 to 190 mm. Methods: We established stand chronologies for total ring width, earlywood and latewood width for the 33 studied stands (985 trees in total). Responses to climate were analysed using pointer years and bootstrapped response functions. Principal component analysis was applied to pointer years and response function coefficients in order to elucidate the ecological structure of the studied stands. Results: Extreme winter frosts are responsible for greater growth reductions in silver fir than in Norway spruce, especially at the upper elevation, while Scots pine was the least sensitive species. Exceptional spring droughts caused a notable growth decrease, especially when local conditions were dry (altitude<1000 m and SWC<100 mm for silver fir, western lowlands for Scots pine). Earlywood of silver fir depended on previous September and November and current‐year February temperature, after which current June and July water supply influenced latewood. Earlywood of Norway spruce was influenced by previous September temperature, after which current spring and summer droughts influenced both ring components. In Scots pine, earlywood and latewood depended on the current summer water balance. Local conditions mainly modulated latewood formation. Conclusions: If the climate becomes drier, low‐elevation dry stands or trees growing in western lowlands may face problems, as their growth is highly dependent on soil moisture availability.     

Xylem sap as a pathway for total mercury and methylmercury transport from soils to tree canopy in the boreal forest

Conifer needles are an important link in the cycling of Total Mercury (THg) and Methylmercury (MeHg) in the boreal ecosystem due to the high THg and MeHg concentrations in litterfall. Translocation within the tree of Hg from soils to the crown canopy has been assumed to be a minor source of the Hg in litterfall. This paper, however, is the first to present direct observations of THg/MeHg transport from the soil via xylem sap. Xylem sap concentrations of THg and MeHg were measured in sap drained from different levels along the boles of freshly cut 100 year old Norway spruce (Picea abies) and Scots pine (Pinus sylvestris). The trees came from a mixed stand growing on podzolized till soils at the Svartberget Forest Research Station in N. Sweden. Soil solution concentrations of THg and MeHg at different levels in the soil profile were measured for comparison.Concentrations of THg in xylem sap ranged from 10–15 ng L^-1 in both the Scots pine and Norway spruce. Concentrations of MeHg varied from 0.03 ng L^-1to 0.16 ng L^-1, with higher values in Scots pine than Norway spruce. If these concentrations are representative of the transport from soils to needles in xylem sap at this site, then only 3% of the MeHg in litterfall (0.12 mg ha^-1 yr^-1) and 11% of the THg (26 mg ha^-1 yr^-1) can originate via this pathway. The upward transport via xylem sap is larger relative to the open field inputs (84% of THg and 17% of MeHg). Comparison of soil solution and xylem sap THg/MeHg suggested some degree of THg exclusion during water uptake in Scots pine and Norway spruce, but MeHg exclusion only in Norway spruce.     

Triacylglycerols in the seeds of northern Scots pine,Pinus sylvestris L., and Norway spruce,Picea abies (L.) Karst.

The geographical variation in the composition of triacylglycerols in seeds of Scots pine (Pinus sylvestris L.) and Norway spruce [Picea abies (L.) Karst.], grown in Finland, was analyzed. The total lipid content of pine seeds was slightly higher in the northernmost provenance (68 °50N), whereas the lipid content of spruce seeds was not affected systematically by the geographical origin of the seeds. The species studied differed in the proportions of fatty acids in their triacylglycerols, though the three most abundant components were the same, i.e. oleic acid (181 n9), linoleic acid (182n-6) and 5,9,12-octadecatrienoic acid (183 5c9c12c). These fatty acids corresponded to more than 80 mol% of the total fatty acids. According to mass spectrometric analyses, the triacylglycerols of both spruce and pine seeds consisted of the same molecular species with 52–56 acyl carbons, but in different proportions. Molecules with 54 acyl carbons represented approximately 75% of the pine and 85% of the spruce triacylglycerols, with the most abundant molecular species being 545, 546 and 547. Some minor differences in the fatty acid composition of triacylglycerols of pine seeds from different seed collecting areas were found: the proportion of linoleic acid slightly increased whereas that of 5,11,14-eicosatrienoic acid decreased towards the northern origins. Similar differences were not found in the proportions of fatty acids in spruce seed lots. Furthermore, the proportions of triacylglycerols in both pine and spruce seeds from northern and southern collecting areas were not significantly different. The higher content of total lipids in spruce seeds compared with pine seeds may be due to the structure of the seed coat, and the lipophilic layers inside it, acting as a barrier to imbibition.     

Effect of open-air fumigation with sulphur dioxide and ozone on phyllosphere and endophytic fungi of conifer needles

The phyllosphere microbial populations inhabiting the needles of three conifer species, Scots pine (Pinus sylvestris L.), Sitka spruce (Picea sitchensis L.) and Norway spruce (Picea abies (L.) Karst.), exposed to SO₂ and O₃, in an open-air fumigation experiment were analysed over a 3 year period using serial dilution after washing, direct plating and a fluorescein diacetate (FDA) enzyme assay. Total fungal populations ranged from 10² to 10⁵ colonyforming units (CPU) g^?1 fresh weight of needles. The dominant fungi isolated from needles varied with tree species and isolation technique; Aureobasidium pullulans (de Bary) Arnaud was most common on Scots pine and Norway spruce and white yeasts on Sitka spruce using the dilution plating method. However, direct plating of needle segments onto culture media indicated that Sclerophoma pythiophila (Corda) Hohnel was dominant on Scots pine and A. pullulans on Sitka and Norway spruce. Green needles of Sitka spruce were found to be endophytically colonized by Rhizosphaera kalkhoffii Bubak, but seldom by Lophodermium piceae (Fuckel) Hohn during extensive sampling in 1990. Statistical analyses revealed significant differences (P<0.05) between plots in the 3 year mean of the total fungal populations or the fungal biomass (FDA assay) on all three tree species. Differences between plots were also observed for a number of dominant component species. Data were also analysed for treatment effects. A significant effect of SO₂ treatment was observed on the total fungal populations on Sitka spruce (P<0.05) which were reduced markedly by the low-SO₂ treatment, while the O₃ treatment caused a significant increase in total fungal numbers on Scots pine (P<0.05). The FDA activity on needles of both Scots pine and Sitka spruce was noticeably higher in the 0₃-only treatment plot, but the overall O₃ effect was not significant. Treatment effects were also detected on the occurrence of component species. The serial dilution method revealed an SO₂ effect (P<0.05) of a reduction in the occurrence of pink yeasts on Sitka spruce and an O₃ effect (P<0.05) of an increase in the occurrence of S. pythiophila on Sitka spruce (P<0.01) but a decrease of Epicoccum nigrum Link and Cladosporium spp. on Scots pine. The direct-plating method revealed an SO₂ effect of an increase in S. pythiophila on Norway spruce (P<0.05). Ozone treatment caused a significant increase in the isolation of a black strain of A. pullulans on Norway spruce (P<0.05). Endophytic colonization of Sitka spruce needles by R. kalkhoffii was found to be increased on two occasions by O₃ exposure.