Relationships between nutrient supply, nitrogen partitioning and growth in young Sitka spruce (Picea sitchensis)

Budget studies have shown that internal cycling may contribute a large proportion of the annual nutrient supply required to support new growth in trees. Use of budgets to quantify internal cycling only quantifies the net transfer of nutrients within the plant. Differential partitioning of remobilized nutrients and current nutrient uptake could lead to errors in the interpretation of results from these studies. We have quantified the dynamic relationships among tree growth, nutrient uptake and internal cycling by labeling the current uptake of N in trees that received contrasting amounts of nutrient. Two-year-old seedlings of Sitka spruce (Picea sitchensis (Bong.) Carr.) were grown in sand culture in a greenhouse for one year. The trees received nutrients in a balanced solution at either a high (high-RAR) or a low (low-RAR) relative addition rate throughout the experiment. Current N uptake was labeled with (15)N from April 13 to July 25. Thereafter, trees were re-potted in clean sand and unlabeled N applied until November 13. Overall growth was sustained for approximately 10 weeks before treatment effects were observed. Initially, no differences in the partition of growth or remobilized N occurred, although partition of current uptake favored the roots of plants in the low-RAR treatment. After 6 weeks, the partition of both growth and remobilized N altered in favor of roots of plants in the low-RAR treatment. Nutrient supply had no effect on the amount or rate of N remobilization. No evidence was found to suggest that N taken up in the current season and partitioned to preexisting shoots or roots is remobilized late in the season to support growth of new shoots. However, some trees in the high-RAR treatment exhibited a second flush of growth later in the season that was partially sustained by remobilization of (15)N from current shoots formed earlier in the season. Use of (15)N demonstrated differential partitioning of current uptake and remobilized N. The results highlight the limitations of simple budget studies for quantifying internal cycling.     

Effects of elevated carbon dioxide and drought on the growth and physiology of clonal Sitka spruce plants (Picea sitchensis (Bong.) Carr.)

Two-year-old Sitka spruce (Picea sitchensis (Bong.) Carr.) plants from four clones were grown in naturally lit growth chambers for 6 months at either ambient (350 ppm) or ambient + 250 ppm (600 ppm) CO(2) concentration. Plants were grown in large boxes filled with peat, in a system that allowed the roots of individual plants to be harvested easily at the end of the growing season. Half of the boxes were kept well watered and half were allowed to dry out slowly over the summer. Plants growing in elevated CO(2) showed a 6.9% increase in mean relative growth rate compared to controls in the drought treatment and a 9.8% increase compared to controls in the well-watered treatment, though there was considerable variation in response among the different clones and water treatments. Rates of net CO(2) assimilation were higher and stomatal conductances were lower in plants grown in elevated CO(2) than in ambient CO(2) in both the well-watered and drought treatments. Both of these factors contributed to the doubling of instantaneous water use efficiency. The partitioning of biomass to roots was unaffected by elevated CO(2), but the ratio of needle mass/stems + branches mass decreased. Together with reduced stomatal conductance, this probably caused the observed increases in xylem pressure potentials with elevated CO(2).     

Needle growth in Sitka spruce (Picea sitchensis): effects of nutrient deficiency and needle position within shoots

Needle development and shoot growth were studied in 14- and 20-year-old Sitka spruce (Picea sitchensis (Bong.) Carr.) planted on an oligotrophic peat and fertilized with N, P and K (control), N and P (-K), N and K (-P) or unfertilized (-NPK). Shoot extension, needle number, and the size and number of needle cells were observed throughout the season. Beginning with bud burst in early May, needle growth continued for nine weeks in the control treatment. Needle growth in the mineral deficiency treatments terminated at the same time as in the controls, although it did not begin until up to three weeks later than in the control trees. Needles developed acropetally along the shoot, with basal needles completing their development first. Cell division in needles of control trees lasted about 3 weeks, with cell expansion continuing for a further 3 weeks. Initial mean cell cycle times were shortest for proximal needles following bud burst, but lengthened as the season progressed. Cell number increased from 6600 in the primordia to 200 000 in mature needles. Final needle dimensions and cell number varied according to the position of the needle on the shoot. The largest needles with the most cells were near the middle of the shoot. Relative to the controls, all mineral deficiency treatments reduced shoot length, bud and needle dimensions, and needle cell number, particularly the -P treatment. Potassium and P deficiency treatments reduced primordial cell number by up to 42%, whereas final mean cell size was 30% greater in the -P treatment, 17% greater in the -NPK treatment, but 14% smaller in the -K treatment. Mean cell size was constant for needles at all positions in any particular treatment at any time, so that final needle size was determined by cell number alone. Needle dry weight/fresh weight ratio continued to increase until early August, with significantly higher ratios in the -K and -P treatments than in the controls.     

Online analysis of volatile organic compound emissions from Sitka spruce (Picea sitchensis)

Volatile organic compound (VOC) emissions from Sitka spruce (Picea sitchensis Bong.) growing in a range of controlled light and temperature regimes were monitored online with a proton transfer reaction-mass spectrometer (PTR-MS) operating at a temporal resolution of approximately 1 min. Isoprene emissions accounted for an average of more than 70% of measured VOCs and up to 3.5% of assimilated carbon. Emission rates (E) for isoprene correlated closely with photosynthetic photon flux (PPF) and temperature, showing saturation at a PPF of between 300 and 400 micromol m(-2) s(-1) and a maximum between 35 and 38 degrees C. Under standard conditions of 30 degrees C and 1000 micromol m(-2) s(-1) PPF, the mean isoprene E was 13 microg gdm(-1) h(-1), considerably higher than previously observed in this species. Mean E for acetaldehyde, methanol and monoterpenes at 30 degrees C were 0.37, 0.78 and 2.97 microg gdm(-1) h(-1), respectively. In response to a sudden light to dark transition, isoprene E decreased exponentially by > 98% over about 3 h; however, during the first 7 min, this otherwise steady decay was temporarily but immediately depressed to approximately 40% of the pre-darkness rate, before rallying during the following 7 min to rejoin the general downward trajectory of the exponential decay. The sudden sharp fall in isoprene E was mirrored by a burst in acetaldehyde E. The acetaldehyde E maximum coincided with the isoprene E minimum (7 min post-illumination), and ceased when isoprene emissions resumed their exponential decay. The causes of, and linkages between, these phenomena were investigated.     

The variation of the basic density within the juvenile wood of Sitka spruce (Picea sitchensis)

The variation in the basic‐density level of 45 stems of Sitka spruce from two Danish sites has been examined. It is shown that the basic‐density level, i.e. the basic density as a function of ring width, within the juvenile wood decreases with (1) increasing distance from the pith and (2) increasing height in the stem (same ring number from pith provided). Furthermore, the basic‐density level seems to decrease with increasing stand quality, which may affect the pattern of variation in basic‐density level if the stand quality class varies with age. The basic density of the juvenile wood is markedly higher in Sitka spruce than in Norway spruce, whereby the risk of stem cracks probably is reduced.     

Branch thickness in unthinned stands of Sitka spruce (Picea sitchensis (Bong.) Carr.)

Results of branch measurings in unthinned stands of Sitka spruce in Denmark are presented. Three different measures of branch thickness are discussed in relation to visual grading of structural timber. On basis of the average thickest branch 5.0 m above ground it is recommended not to let spacing exceed 1.8x1.8 m‐2.0x2.0 m at establishment. Since Sitka spruce branches react profusely when spacing widens, this is the upper limit, presupposing a no‐thinning regime. With reference to the literature, the same recommendation can be given on basis of machine stress gradings.     

Resistance to the green spruce aphid (Elatobium abietinum Walker) in progenies of Sitka spruce (Picea sitchensis (Bong) Carr.)

Damage caused by Elatobium abietinum was estimated as needle loss in a progeny trial of Sitka spruce in Denmark in 1990 and 1992. The parents of the progeny had been selected for putative aphid resistance during a massive attack of aphids in 1957. Needle loss was significantly less in the progeny than in a standard provenance. Family heritability was estimated as 0.72 (P>0.0001). Aphid damage had an impact on growth and vigour of trees following warm winters in 1989 and since, but no correlation could be found between growth parameters and damage of different families. Due to high heritability and large variation, high gains are expected by breeding. Practical and theoretical consequences of including aphid resistance in breeding programmes of Sitka spruce are discussed.     

Prediction of transverse shrinkages of young-growth Sitka spruce (Picea sitchensis) and western hemlock (Tsuga heterophylla) with ultrasonic measurements

Abstract

The objective of this study was to examine the potential of acoustic measurement as a rapid and nondestructive method to predict the dimensional stability of young-growth Sitka spruce and western hemlock. Ultrasonic velocity, peak energy, specific gravity, and radial and tangential shrinkages were measured on twenty-four 25-×102-×y 25-mm specimens obtained from a 58-year-old stand in Southeast Alaska. We found that specific gravity and peak energy of ultrasonic signals were not good predictors of transverse shrinkage, as indicated by poor correlations. Ultrasonic velocity, on the other hand, was found to be a significant predictor of transverse shrinkage and therefore has good potential to be used as a field method to evaluate dimensional stability. The single-parameter prediction model explained 86% of transverse shrinkages in western hemlock and 71% of transverse shrinkages in Sitka spruce. Further study is needed to test the capability and feasibility of using acoustic velocity to predict both longitudinal and transverse shrinkages of wood in standing trees.     

The effect of seed pretreatment and sowing date on the nursery emergence of Sitka spruce (Picea sitchensis [Bong.] Carr.) seedlings

Sitka spruce seeds which had been: (1) untreated (U), (2) prechilled(P), or (3) prechilled and then redried and stored for 16 months(R), were sown into progressively warmer seed beds on four datesat monthly intervals from March to June. At each sowing date,samples of seed were germinated in the laboratory at optimum(20°C) and sub-optimum (10°C) temperatures. Prechillingand prechilling + redrying both increased the rate and totalamount of germination compared with untreated seed at each testtemperature. However in the nursery, the beneficial effectsof pretreating Sitka spruce seed were restricted to sowing inMarch and April when soil temperatures were sub-optimal forthe germination of untreated seeds. Pretreatment resulted inrapid seedling emergence at soil temperatures between 10°Cand 15°C, whereas untreated seed germinated slowly and seedlingemergence continued until high soil temperatures in June killedungerminated seeds. The pattern of seedling emergence from theredried seed was similar to that from the prechilled seed, exceptfor a lag in the start of seedling emergence probably due tothe extra time needed for redried seed to imbibe water in theseed beds. Thus redrying caused insignificant loss of seed performanceon the nursery. Seed sown in May and June experienced average temperatures closeto the optimum for germination of 20°C, however daily maximaregularly exceeded 35°C resulting in the cessation of seedlingemergence in all three seed treatments, and the loss of newlyemerged seedlings. Periods of warm weather from June onwardswere the main factors limiting the final number and size ofseedlings.     

Long-term effects of elevated carbon dioxide concentration and provenance on four clones of Sitka spruce (Picea sitchensis). II. Photosynthetic capacity and nitrogen use efficiency

Four clones of Sitka spruce (Picea sitchensis (Bong.) Carr.) from two provenances, at 53.2 degrees N (Skidegate a and Skidegate b) and at 41.3 degrees N (North Bend a and North Bend b), were grown for three growing seasons in ambient (~350 micromol per mol) and elevated (~700 micromol per mol) CO2 concentrations. The clones were grown in stress-free conditions (adequate nutrition and water) to assess the effect of elevated [CO2] on tree physiology. Growth in elevated [CO2] significantly increased instantaneous photosynthetic rates of the clonal Sitka spruce saplings by about 62%. Downward acclimation of photosynthesis (A) was found in all four clones grown in elevated [CO2]. Rubisco activity and total chlorophyll concentration were also significantly reduced in elevated [CO2]. Provenance did not influence photosynthetic capacity. Best-fit estimates of Jmax (maximum rate of electron transport), Vcmax (RuBP-saturated rate of Rubisco) and Amax (maximum rate of assimilation) were derived from responses of A to intercellular [CO2] by using the model of Farquhar et al. (1980). At any leaf N concentration, the photosynthetic parameters were reduced by growth in elevated [CO2]. However, the ratio between Jmax and Vcmax was unaffected by CO2 growth concentration, indicating a tight coordination in the allocation of N between thylakoid and soluble proteins. In elevated [CO2], the more southerly clones had a higher initial N use efficiency (more carbon assimilated per unit of leaf N) than the more northerly clones, so that they had more N available for those processes or organs that were most limiting to growth at a particular time. This may explain the initial higher growth stimulation by elevated [CO2] in the North Bend clones than in the Skidegate clones.     

Long-term effects of elevated carbon dioxide concentration and provenance on four clones of Sitka spruce (Picea sitchensis). I. Plant growth, allocation and ontogeny

Four clones of Sitka spruce (Picea sitchensis (Bong.) Carr.) from two provenances, at 53.2 degrees N (Skidegate a and Skidegate b) and at 41.3 degrees N (North Bend a and North Bend b), were grown near Edinburgh (55.5 degrees N), U.K., for three growing seasons in ambient (~350 micromol per mol) and elevated (~700 micromol per mol) CO2 concentrations under conditions of non-limiting water and nutrient supply. Bud phenology was not affected by elevated [CO2] in the second growing season, but in the third year, the duration of shoot extension growth in three of the four clones (North Bend clones and Skidegate a) was significantly shortened, because of the suppression of lammas growth. Saplings in elevated [CO2] had significantly greater dry masses of all components than saplings in ambient [CO2]. However, comparison of relative component dry masses between plants of similar size showed no effect of [CO2] treatment on plant allometric relationships. This finding, and the observed suppression of lammas growth by high [CO2] during the third growing season suggests that the main effect of increasing [CO2] is to accelerate sapling development. Clonal provenance did not affect dry mass production in ambient [CO2]. However, in elevated [CO2] the more southerly clones significantly out-performed the more northerly clones when grown at a latitude close to the latitudinal provenance of the Skidegate clones. As atmospheric carbon dioxide concentration rises, such changes in the relative performance of genotypes may be exploited for economic gain through appropriate selection of provenances for forest plantings.     

Effects of silicon-rich soil amendments on growth,mortality and bark feeding damage of Sitka spruce (Picea sitchensis) seedlings under field conditions

Alternative methods of protection are required against feeding by the large pine weevil (Hylobius abietis) on the bark of conifer seedlings. Silicon (Si) has been shown to enhance the resistance of plants to insect herbivores. This study investigated the effects of low doses of Si-rich soil amendments on growth, mortality and bark feeding damage of Sitka spruce (Picea sitchensis) seedlings. Two-year old seedlings were grown, individually, in soil taken from a tree nursery treated with coal ash, peat ash, rice husk ash, slag, sodium metasilicate or a commercially available Si fertiliser (Pro-Tekt) and planted out on two reforestation sites in Ireland. Seedlings grew well (about 20% growth in terms of height, 66% in root collar diameter, after two growing seasons), and Si-rich amendments did not have a significant effect on growth or mortality. Bark feeding damage on Si-treated seedlings did not vary significantly from control seedlings. Bark Si concentrations were not significantly larger in treated seedlings than in control seedlings, but control seedlings already had comparatively high bark Si concentrations (560?mg?kg^?1 dry tissue). In conclusion, Sitka spruce seedlings grown in the presence of Si-rich soil amendments prior to planting did not show greater resistance to weevil feeding under the present conditions.     

Variation in the levels of the antifungal stilbene glucosides astringin and isorhapontin in the bark of Sitka spruce [Picea sitchensis (Bong.) Carr.

The concentrations of the constitutive antifungal stilbene glucosides astringin and isorhapontin were measured in the bark of a number of Sitka spruce [Picea sitcbensis (Bong.) Carr.] genotypes. These included trees of different provenances, trees grown at different sites, and young clonal plants. The stilbene contents of individual trees or clones varied widely. In general this was attributable to variation between individual genotypes, and was not related to provenance or site.     

Physical and chemical responses of Sitka spruce (Picea sitchensis) clones to colonization by Heterobasidion annosum as potential markers for relative host susceptibility

Forty-one 2-year-old clones of Picea sitchensis (Bong.) Carr. from three full-sib families (14 clones from each of two families and 13 clones from a third family) were either wounded and inoculated with an isolate of Heterobasidion annosum (Fr.) Bref. or wounded without inoculation. Lesion lengths on the inner bark from the point of inoculation varied among clones 35 days after treatment. There was no relationship between lesion length and relatedness within families. Two clones (21342 and 25202) with the shortest lesions, tentatively designated as less susceptible to H. annosum, and two clones (21176 and 27166) with the longest lesions, designated more susceptible, were selected for comparison of host anatomical and chemical responses to infection. The position and structure of the ligno-suberized boundary zone (LSZ) in the bark of the clones suggested that the less susceptible clones formed thicker layers of suberized cells in the LSZ following wounding plus inoculation. No LSZ was observed in two ramets of the more susceptible Clone 27166 following wounding and inoculation with H. annosum. Compared with more susceptible genotypes, clones of P. sitchensis with low susceptibility to H. annosum had high relative proportions of (+)-alpha-pinene, (-)-beta-pinene and one unidentified terpene constituent (Unknown-15) in cortical resin sampled 25 cm from the lesions. In contrast, more susceptible clones had higher relative proportions of (-)-limonene, Unknown-16, Unknown-18 and Unknown-19. In the secondary resin produced in bark tissues surrounding the lesions, proportions of several monoterpene constituents varied; these changes included a decrease in the relative amount of beta-phellandrene and corresponding small increases in some minor constituents. The concentrations of the monoterpenes, except a few minor constituents, increased in the infected tissues. Wounding plus inoculation with H. annosum resulted in varied monoterpene responses, with distinct differences between less susceptible and more susceptible clones. In less susceptible clones, Unknown-19 increased following wounding plus inoculation, whereas in more susceptible clones, concentrations of delta-3-carene and Unknown-13 and Unknown-16 increased. Differences in both constitutive and induced resin monoterpene profiles may provide useful markers for resistance to H. annosum in selection and breeding programs.     

An outbreak of Vapourer Moth (Orgyiaantiqua L.: Lepidoptera Lymantridae) on Sitka spruce (Picea sitchensis (Bong.) Carr.) in Central Scotland

Highly significant differences were observed in the intensityof defoliation on Sitka spruce of different heights at CarronValley Forest, Central Region. Most severe defoliation occurredon the smallest trees. Significant differences also occurredbetween degree of defoliation and foliar potassium and phosphorusconcentrations. Winter and summer rainfall patterns immediatelyprior to the outbreak showed higher than average winter andlower than average summer rainfall compared with the 30-yearmean. Experience with this pest suggests that it will continueto have local nuisance value.     

Comparison of Crop Rotations, and of Fertilizer with Compost, in long-term Experiments with Sitka spruce (Picea sitchensis) in two English Nurseries

Groups of related experiments, of more than 300 plots, withseedlings and transplants of Picea sitchensis, tested inorganicfertilizer and bracken-hopwaste compost, and compared continuousconifer cropping with a rotation in which one conifer crop inthree was replaced by fallow or a green manure crop. The experiments,lasting 15 years, were done in a nursery on moderately acidagricultural land at Kennington and in a heathland nursery onan acid sandy podsol at Wareham. At Kennington, seedlings given fertilizer grew better throughoutthan those given compost, with no additional benefit when bothwere given, despite considerable increases in soil organic matterand nutrients on compost-treated plots. On plots with fertilizer,growth was maintained during the experimental period. At Wareham,fertilizer lost its initial advantage; neither of the manunaltreatments supplied adequate amounts of major nutrients butfertilizer and compost together was better than either alone. With transplants, at both nurseries, fertilizer was slightlymore effective during the early years, but compost later. Interrupting continuous conifer cropping by fallow or greencrops had no advantage at either nursery.