SURFACE WAX DEPOSITS ON FOLIAGE OF PICEA PUNGENS AND OTHER CONIFERS

Epicuticular waxes of five cultivated varieties of Picea pungens and six other species of conifers were examined directly with a scanning electron microprobe. The quality, quantity, and distribution patterns of surface wax deposits varied between species, within species, within plants, and within individual leaves. Two basic types of wax were observed, structural and amorphous. Structural wax is associated with glaucousness and the blue foliage color in Picea pungens and other species. The degree of occlusion of the stomatal pore by surface waxes also varied from heavily occluded to completely free of wax. Wax in the epistomatal chamber may be another resistance barrier to gas exchange by conifer needles.     

The histopathology of Phaeocryptopus gaeumannii on Douglas-fir needles

Germinating ascospores of Phaeocryptopus gaeumannii produce suprastomatal appressoria from which penetration pegs enter needles. Initial infection occurs between late May and early Jul and coincides with budbreak and shoot elongation. Colonization within needles is exclusively intercellular and increases continuously during Jul-May. No intracellular hyphae or haustoria were observed, but hyphae closely appressed to mesophyll and palisade cell walls are abundant by 3-5 mo after initial infection. Pseudothecial primordia begin to form in epistomatal chambers Oct-Apr, 4-9 mo after initial infection. Pseudothecial primordia developing in the epistomatal chamber are connected to the endophytic thallus by specialized cells in the substomatal chamber that have thickened apical walls and resemble phialides but are not involved in asexual reproduction. The apical wall thickenings instead appear to function as reinforcement against the turgor pressure of the guard cells, allowing cytoplasmic continuity to be maintained between the developing pseudothecium and vegetative hyphae within the needle. Concurrent with the formation of pseudothecial primorida, epiphytic hyphae emerge from the periphery of developing pseudothecia, grow across the needle surface, form numerous anastomoses and reenter the needle by producing appressoria above unoccupied stomata. Epiphytic hyphae and their associated appressoria gradually become more abundant during Oct-Jan.     

The Loss of Structural Integrity in Damaged Spruce Needles from Locations Exposed to Air Pollution

Abstract Within the framework of appraising the new type of forest damage, the epidermis of two-year-old spruce ( Picea abies ) needles was investigated as a possible site for primary attack by air pollutants using light and scanning electron microscopy.
Depending upon the extent of the macroscopically visible needle leaf disease, a degeneration process of the cuticular, stomatal and epidermal structures leading to the lethal destruction of the needle surface was observed.
Stomatal and epidermal cells become increasingly deformed or collapsed showing increasing discolouration due to phenolic deposits. Additionally, the erosion of stomatal waxes is accelerated, leading initially to a temporary occlusion of the stomata, and then to a wide opening of the antechamber following a complete decomposition of the plug.
Surface lesions indicating early damage to the epidermal tissue already occur on slightly yellowed needles. The barrier effect of the spruce needle epidermis on exogenous stressors could thereby already be reduced at an early stage, triggering incipient losses of structural resistance.     

Morphological and stomatal responses of Norway spruce foliage to irradiance within a canopy depending on shoot age

Morphological and stomatal responses of Norway spruce (Picea abies) foliage to light availability were studied in respect to shoot age. Needle minor diameter (D(1), anatomical width), major diameter (D(2), anatomical thickness), dry weight (M), and tissue density index (I(D)) increased, and needle flatness (Fl) and specific leaf area (SLA) decreased with foliage age, while shade foliage demonstrated higher morphological plasticity as compared to sun foliage. Needle minor diameter, dry weight, and the ratio of total to projected leaf area increased, and needle flatness and specific leaf area decreased with daily average photosynthetic photon flux density (Q(D)). The current-year foliage exhibited the highest variation with irradiance, while the morphological plasticity decreased with needle ageing. The morphological characteristics of needles were independent of irradiance if Q(D) was above 300 μmol m(-2) s(-1). D(1) was the only linear needle characteristic which significantly changed with light availability within a canopy, and thus determined needle flatness, SLA, as well as the ratio of total to projected leaf area (TLA/PLA). Needle flatness was a characteristic responding most sensitively to the photosynthetic photon flux density, R(2) was 0.68, 0.44, and 0.49 for the current-year, 1-year-old, and 2-year-old foliage, respectively. TLA/PLA ranged from 2.2 to 4.0 depending on D(1). Variation in SLA in response to light availability can be attributed to changes both in needle shape and tissue density. Stomatal responses to photosynthetic photon flux density (Q(P)) depended on foliage type (sun or shade) and age. Sun needles demonstrated higher daily maximum leaf conductances to water vapour compared to shade needles. The shade needles responded more sensitively to changes in Q(P) at dawn and sunset than the sun needles, while older needles of both foliage types exhibited faster stomatal responses. The light-saturation of leaf conductance (g(L)) was achieved by 20 μmol m(-2) s(-1) for shade foliage, and approximately by 50 μmol m(-2) s(-1) for sun foliage. As a rule, g(L) changed in response to irradiance faster in the evening, i.e. at decreasing irradiance. Stomata were not usually completely closed in the dark before sunrise and after sunset, the phenomenon being more pronounced in older shoots and sun needles. Nightly water losses from spruce foliage are attributable primarily to older shoots, and are related to age-dependent changes in stomatal responsiveness.     

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.     

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.     

Diversity of endophytic fungi of single Norway spruce needles and their role as pioneer decomposers

The diversity of endophytic fungi within single symptomless Norway spruce needles is described and their possible role as pioneer decomposers after needle detachment is investigated. The majority (90%) of all 182 isolates from green intact needles were identified as Lophodermium piceae. Up to 34 isolates were obtained from single needles. Generally, all isolates within single needles had distinct randomly amplified microsatellite (RAMS) patterns. Single trees may thus contain a higher number of L. piceae individuals than the number of their needles. To investigate the ability of needle endophytes to act as pioneer decomposers, surface-sterilized needles were incubated on sterile sand inoculated with autoclaved or live spruce forest humus layer. The dry weight loss of 13-17% found in needles after a 20-week incubation did not significantly differ between the sterilized and live treatments. Hence, fungi surviving the surface sterilization of needles can act as pioneer decomposers. A considerable portion of the needles remained green during the incubation. Brown and black needles, in which the weight loss had presumably taken place, were invaded throughout by single haplotypes different from L. piceae. Instead, Tiarasporella parca, a less common needle endophyte, occurred among these invaders of brown needles. Needle endophytes of Norway spruce seem thus to have different abilities to decompose host tissues after needle cast. L. piceae is obviously not an important pioneer decomposer of Norway spruce needles. The diversity of fungal individuals drops sharply when needles start to decompose. Thus, in single needles the decomposing mycota is considerably less diverse than the endophytic mycota.     

Penetration of blue nad UV radiation measured by fiber optics in spruce and fir needles

The penetration of blue light (460 nm) and UV-A (360 nm) radiation into needles of spruce ( Picea engelmannii Parry ex. Engelm.) and fir [ Abies lasiocarpa (Hook.) Nutt. var. Iasiocrpa ] was examined. The distribution of collimated and scattered light in intact needles was measured by quartz fiber optic microprobes. Both blue light and UV-A were attenuated more rapidly with depth in fir than in spruce, although the light profiles for the two light regimes were different; blue light penetrated to a greater depth than UV-A, Removal of the epicuticular wax from the needle surface increased the amount of internal blue light slightly, but had little effect on the amount of UV-A. The differences in light penetration in light penetration in spruce and fir may be caused bhy different pigmentation and leaf anatomy.     

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.     

Monoterpenes in sitka spruce: Within tree and seasonal variation

The composition of oils from needles and cortex of Sitka spruce is unique. Cortex oil is essentially all monoterpene hydrocarbons, while needle oil may be up to 50% oxygenated monoterpenes. Very wide seasonal variations in composition occur in needle oil in young tissue. At bud burst, the oil is > 95% myrcene; this drops to about 40% at the end of summer. The oxygenated terpenes camphor and piperitone develop to about 20% each in concentration during the growing season.     

Photosynthetic action spectra of trees: I. Comparative photosynthetic action spectra of one deciduous and four coniferous tree species as related to photorespiration and pigment complements

Comparative isoenergetic action spectra of net photosynthesis for intact, current year foliage of five tree species were determined from 400 to 710 nm by CO₂ exchange analysis. The blue (400 to 500 nm) peak of net photosynthetic activity for the green broadleaves of red alder (Alnus rubra Bong.) was reduced to a plateau for the green needle-leaves of Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco) and Sitka spruce (Picea sitchensis [Bong.] Carr.), a shoulder for the blue-green needles of Colorado spruce (Picea pungens Engelm.), and a reduced shoulder for the blue-white needles of Blue spruce (Picea pungens var. hoospii). These differences were attributable neither to a differential blue light stimulation of photorespiration nor to a differential presence of a nonplastid screening pigment. The conifers all had similar carotenoid-chlorophyll ratios, with approximately 50% more carotenoid relative to chlorophyll as compared to red alder. Blue light absorption and low efficiency of energy transfer by the carotenoids probably accounts for the low net photosynthetic activity of the green conifers in blue light as compared to red alder. Leaf form per se (broad versus needle) had no distinguishable influence on these results.     

Estimating the needle area from geometric measurements: application of different calculation methods to Norway spruce

Different calculation methods, based on needle geometry, for estimating both projected area (PLA) and total surface area (TLA) of foliage in Norway spruce [Picea abies (L.) Karst.] were compared. Seventy-eight shoots of four age classes were sampled from both the basal and top thirds of crowns. Three dimensions (the length, minor and major diameters) of needles were taken, and the needle shape was approximated to a parallelepiped or ellipsoid. There was a perfect coincidence of the measured and estimated values of PLA calculated as the width of the needle projection multiplied by needle length, and corrected for needle taper (method III), or when the needle projection was treated as a rectangle joined with half-ellipses at both ends (method IV). The most reliable estimations of TLA resulted from treating the needle sides as faces of the parallelepiped tapering at their ends in the form of half-ellipses. The ratio of TLA to PLA ranged from 2.2 to 4.0 depending on the needle morphology. Needle minor diameter (anatomical width; D ₁) was found to be a better morphological index of the spruce foliage than needle flatness, i.e. the ratio of major to minor diameter. Expressing the factor for converting PLA to TLA as a function of D ₁ considerably improved the precision of the estimates. Close relationships were established between specific leaf area, expressed on both a projected area (SLA_P) and total surface area basis (SLA_T), needle dry weight (R ² was 0.799 and 0.852, respectively) and minor diameter of needles (R ² was 0.701 and 0.554, respectively). Received: 14 April 1998 / Accepted: 23 September 1999     

Is long-lived foliage in Picea mariana an adaptation to nutrient-poor conditions?

Summary This study evaluated the contribution of different ages of foliage to the nutrient and carbon balance of black spruce (Picea mariana (Mill.) B.S.P.) from a nutrient-poor peatland in Alberta. Seasonal patterns of foliar nitrogen and phosphorus concentration and content were examined in six needle cohorts up to 10 years old. Trees were treated to simulate excess nutrient deficiency (removal of all one-year-old foliage), nutrient excess (fertilized with 250, 50, 100 kg ha^–1 NPK split application in June and July), or left as controls. Gas exchange (net assimilation-Na, stomatal conductance-g_s, mesophyll conductance-g_m, water-use efficiency-WUE, dark respiration-RS) was measured on six different needle cohorts in several control trees in 1989 and 1990. Nitrogen and phosphorus concentration decreased with needle age. Foliar nutrient concentration fell from April to June and then was stable until September except for the fertilized trees where it increased. There was no evidence of greater than normal retranslocation of nutrients from older needles for defoliated trees or greater than normal nutrient loading in older needles of fertilized trees. NA, g_s, g_m, WUE, and RS were similar for all needles up to six or eight years old, these older needles having NA of 65% of current needles and similar RS. The results do not support to conclusion that older needles of black spruce are retained as an adaptation to nutrient stress. It does not appear that older needles serve as a nutrient storage site in conditions of excess nutrient availability or a greater than normal nutrient source during times of excess nutrient deficiency. It appears that the maintenance of long-livedfoliage in black spruce does not provide for greater flexibility in tree nutrient allocation. Their contribution to the carbon balance of the tree seems to be sufficient to explain their retention.     

The effect of previously infested spruce needles on the growth of the green spruce aphid, Elatobium abietinum, and the effect of the aphid on the amino acid balance of the host plant

The effect of chlorosis induced in needles of Sitka and Norway spruce by the green spruce aphid on growth of the aphid is investigated, and the effect of infestation of the aphid on amino acid levels in Sitka spruce foliage is reported. On both Sitka and Norway spruce green spruce aphids were heavier when reared on chlorotic (previously infested) needles than when reared on green (previously uninfested) needles. The effect was more pronounced on Sitka than on Norway spruce. Infestation of the aphid altered the amino acid balance of Sitka spruce foliage but not the concentration of total amino acids. Possible causes of chlorosis, the influence of individual amino acids on aphid growth, the potential effect of chlorosis on outbreaks of the aphid and the differences in susceptibility of Sitka and Norway spruce to damage by the aphid are discussed.     

Ultraviolet-B and visible light penetration into needles of two species of subalpine conifers during foliar development

The depth of penetration of Ultraviolet-B (UV-B, 300 and 320 nm) and visible (680 nm) light was measured in foliage of Abies lasiocarpa and Picea engelmannii using a fibre-optic microprobe. Measurements were made on foliage at four times during development: needles were sampled from within expanding buds (in bud); within 72 h of emergence from the bud scales (emergent); from elongating branches (elongating); and from foliage that emerged the previous summer (mature). Light attenuation in pre-emergent needles of both species was steep and showed strong wavelength dependence. Short wavelength 300-nm light was attenuated strongly in the developing epidermal layer, but a significant proportion of this potentially damaging UV-B radiation penetrated into the mesophyll. For A. lasiocarpa and P. engelmannii, 99% attenuation of 300-nm light occurred at 51 and 96 μm, respectively, well within the mesophyll. At this stage, however, the bud scales were opaque to light below 400nm. As the epidermal cell walls and cuticle continued to develop and chlorophyll accumulated following emergence from the bud scales, light attenuation, particularly of UV-B radiation, increased. Although no UV-B is transmitted through the epidermis-hypodermis of mature needles, small but measurable quantities of 300- and 320-nm light were measured in the photosynthetic mesophyll of post-emergent and elongating needles. Thus, shortly after emergence from the bud scales in mid-June to mid-July, when incident UV doses are highest, absorption of UV-B radiation by potentially sensitive chromophores in the mesophyll may disrupt physiological and developmental processes in these species. Soluble UV-absorbing pigments accumulated during needle maturation for P. engelmannii but not A. lasiocarpa, suggesting that, for A. lasiocarpa at least, the development of effective UV screening properties in the epidermis may not be related to the induction of soluble flavonoids.     

Damage caused by the Green Spruce Aphid to Norway and Sitka spruce needles

Three-month-old needles of Sitka spruce were less susceptible to Elatobium abietinutn than 15-month-old needles. Symptoms appeared after longer aphid feeding times but only a proportion of damaged needles fell. After short feeding periods symptoms appeared in more Norway spruce needles than in Sitka spruce, whereas longer feeding periods resulted in more needles producing symptoms in the Sitka spruce. The symptoms took 4—6 days longer to appear in Norway spruce, and needle fall followed a longer feeding period than on Sitka spruce. Following 72 h feeding, needle fall occurred more quickly on Sitka spruce than on Norway spruce. The time taken for needle fall to occur was inversely related to the feeding time in Sitka spruce but such a response was not evident in Norway spruce. The results are discussed in relation to the differences exhibited in the probing behaviour of the aphid on the two spruces.     

Ribulose-l,5-bisphosphate carboxylase activity and influence of air pollution in spruce

Methods were established, which render possible a simultaneous determination of ri-bulose-l,5-bisphosphate (RuBP) carboxylase (EC 4.1.1.39) activity and chlorophyll content of Norway spruce (Picea abies Karst.) needles from a detergent-containing aqueous crude extract. Spruce RuBP carboxylase was tentatively characterized with regard to kinetic properties. Recovery experiments employing purified wheat RuBP carboxylase proved quantitative extraction of the enzyme from spruce foliage. Five timber stands consisting of 35–62 years old spruce, two of which exhibited the typical symptoms of recent spruce decline, were compared. For the needle generations 1 to 4 the enzyme activities as well as chlorophyll and protein concentrations were determined. The results do not indicate an involvement of RuBP carboxylase in spruce decline.     

Temperature sensitivity of nitrogen productivity for Scots pine and Norway spruce

Environmental conditions control physiological processes in plants and thus their growth. The predicted global warming is expected to accelerate tree growth. However, the growth response is a complex function of several processes with both direct and indirect effects. To analyse this problem we have used needle nitrogen productivity, which is an aggregate parameter for production of new foliage. Data on needle dry matter, production, and nitrogen content in needles of Scots pine ( Pinus sylvestris) and Norway spruce ( Picea abies) from a wide range of climatic conditions were collected and needle nitrogen productivities, defined as dry matter production of needles per unit of nitrogen in the needle biomass, were calculated. Our results show that the nitrogen productivity for spruce is insensitive to temperature. However, for pine, temperature affects both the magnitude of nitrogen productivity at low needle biomass and the response to self-shading but the temperature response is small at the high end of needle biomass. For practical applications it may be sufficient to use a species-specific nitrogen productivity that is independent of temperature. Because temperature affects tree growth indirectly as well as through soil processes, the effects of temperature change on tree growth and ecosystem carbon storage should mainly be derived from effects on nitrogen availability through changes in nitrogen mineralization. In addition, this paper summarises data on dry matter, production and nitrogen content of needles of conifers along a temperature gradient.     

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.     

Photosynthetic Action Spectra of Trees: II. The Relationship of Cuticle Structure to the Visible and Ultraviolet Spectral Properties of Needles from Four Coniferous Species

The relative reflectance spectra for control and treated (surface wiped) current-year foliage of Douglas fir, and Sitka, Colorado, and Blue spruce (Pseudotsuga menziesii [Mirb.] Franco, Picea sitchensis [Bong.] Carr., Picea pungens Engelm., and Picea pungens Engelm. var. hoopsii, respectively) were obtained from 220 to 700 nm. The green color of the control foliage of both Douglas fir and Sitka spruce was unaffected by the treatment whereas the blue-green and blue-white foliage of control Colorado and Blue spruce, respectively, became “green” as a result of the wiping. The relative reflectance curves for all green foliage, including the treated Colorado and Blue spruce, were all very similar with a peak in the green (540-560 nm), minima in the red (660-680 nm) and blue (450-500 nm), and very low reflectivities in the ultraviolet (λ < 400 nm). In contrast, the control foliage for Colorado and Blue spruce both showed a generally higher relative reflectance over most of the visible spectrum (400-700 nm) with a marked increase in the blue region (400-500 nm). At wavelengths below 420 nm, their relative reflectances increased sharply with decreasing wavelength, the reflectance at 220 nm for Blue spruce being over four times that at 540 nm.