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Sexual reproduction in a greenhouse and reduced autumn frost hardiness of Picea abies progenies.

In 1989, identical crosses (2-3 females within males) were performed with Picea abies (L.) Karst. in a greenhouse seed orchard at Biri nursery and in an outdoor seed orchard at Huse, 32 km north of Biri. Pollination began 17 days earlier in the greenhouse than outdoors at Huse. The potted grafts in the greenhouse were moved outdoors when the seed cones were no longer receptive. Twelve full-sib family pairs (Biri and Huse) from these crosses were grown in a phytotron and tested for height and autumn frost hardiness during their first growing season. No significant difference was found between the indoor (Biri) and outdoor (Huse) progenies for height growth. However, the progenies from the greenhouse seed orchard were significantly more susceptible to frost than their full-sibs from the outdoor seed orchard. There was no significant interaction between males and the flowering environment, but a significant female x flowering environment interaction was present as a result of greater differences in frost hardiness between progenies from females in the greenhouse seed orchard than in the outdoor seed orchard. Although seeds from the outdoor seed orchard generally had a greater biomass than seeds from the greenhouse seed orchard, the difference in seed weight did not explain the difference in frost hardiness. We hypothesize that temperature and photoperiod during pollination and fertilization affect the frost hardiness of the progenies.

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

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

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Functional biodiversity of microbial communities in the rhizospheres of hybrid larch (Larix eurolepis) and Sitka spruce (Picea sitchensis).

The diversity of microorganisms associated with trees and their different functional capabilities is thought to be a consequence of variation in carbon compounds in the rhizosphere. We used the Biolog(R) system (Biolog Inc., Hayward, CA), a redox-based test, to construct sole carbon source utilization profiles (metabolic fingerprints) of microbial communities from the rhizospheres and rhizoplanes of hybrid larch (Larix eurolepis A. Henry) and Sitka spruce (Picea sitchensis Bong. Carr.) taken from a farm woodland site and two second-rotation plantation forest sites. Canonical variate analysis (CVA) of carbon utilization data differentiated among the microbial communities from the three forest sites, with the greatest discrimination between the farm woodland and the two second-rotation forest sites. Carbohydrates and carboxylic acids were the substrates responsible for this discrimination. Carbon profiles of the microbial communities from the rhizospheres of the two tree species also clustered when evaluated by CVA, as a result of differences in utilization of carboxylic acids and amino acids, suggesting that these tree species differ in the exudates they produce. Isolation and enumeration of organisms confirmed that there were qualitative and quantitative differences in the culturable populations of microorganisms at the different sites and between tree species. We conclude that Biolog is a useful technique for evaluating the functional diversity of microbial communities; however, to interpret the results accurately, they must be assessed in conjunction with the actual carbon substrates available in the particular ecosystem under study.

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Responses of Picea, Pinus and Pseudotsuga roots to heterogeneous nutrient distribution in soil.

The spatial distribution of plant-available mineral nutrients in forest soils is often highly heterogeneous. To test the hypothesis that local nutrient enrichment of soil leads to increased root proliferation in the nutrient-rich soil zone, we studied the effects of nutrient enrichment on the growth and nutrient concentrations of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) roots. Three-year-old seedlings were grown for 9 months in split-root containers filled with nutrient-poor forest mineral soil, with one side supplemented with additional mineral nutrients. Root dry weight and root length in Scots pine and Norway spruce were increased in the nutrient-supplemented soil compared with the nonsupplemented side, whereas root growth in Douglas-fir was unaffected by nutrient enrichment. Of the three species examined, Norway spruce exhibited the highest root and shoot growth and the highest nutrient demand. Specific root length (m g(-1)) and the number of root tips per unit root length were not affected by local nutrient addition in any of the species. Despite increased root growth in Norway spruce and Scots pine in nutrient-supplemented soil, their root systems contained similar nutrient concentrations on both sides of the split-root container. Thus, coniferous trees may respond to local nutrient supply by increased root proliferation, but the response varies depending on the species, and may only occur when trees are nutrient deficient. As a response to local nutrient enrichment, increases in root dry matter or root length may be better indicators of pre-existing nutrient deficiencies in conifers than increases in root nutrient concentrations.

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Influences of altered phytohormone use on endogenous ABA and mRNA populations during white spruce (Picea glauca) somatic embryo culture.

We examined endogenous abscisic acid [(+)-ABA] amounts and mRNA populations in white spruce (Picea glauca (Moench) Voss) somatic embryos in liquid suspension culture during a pretreatment prior to induction of maturation with exogenous (+)-ABA. The pretreatment consisted of removal of 2,4-dichlorophenoxyacetic acid (2,4-D) from the multiplication medium for 7 days. The removal of 2,4-D resulted in a slight increase in endogenous ABA in pretreated tissues compared to nonpretreated tissues at the end of the 7-day pretreatment period (42.4 versus 20.0 pg mg(-1) lyophilized tissue). Altered gene expression patterns were observed as early as one day after the start of the pretreatment, with more than 17 new polypeptides found in pretreated tissues. The influence of pretreatment continued to be observed after tissues were transferred to ABA-containing maturation medium. By comparing mRNA populations in pretreated and nonpretreated tissues cultured on either ABA-containing or ABA-free maturation medium, at least 12 mRNAs were observed to be induced by ABA, among which three polypeptides were ABA inducible only in pretreated tissues.

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Acclimation to low irradiance in Picea abies: influences of past and present light climate on foliage structure and function.

Leaf retention time increases with decreasing irradiance, providing an effective way of amortizing the costs of foliage construction over time. To elucidate the physiological mechanisms underlying this dependence, I studied needle life span, morphology, and concentrations of carbon, nitrogen and nonstructural carbohydrates along a gradient of relative irradiance in understory trees of Picea abies (L.) Karst. Maximum needle life span was greater in shaded trees than in sun-exposed trees. However, irrespective of irradiance, needles with maximum longevity were situated in the middle rather than the bottom of the canopy, suggesting that needle life span is determined by the irradiance to which needles are exposed during their primary growth. Morphology and chemistry of current-year needles were adapted to prevailing light conditions. Current-year needles exposed to high irradiances had greater packing of foliar biomass per unit area than shaded needles, whereas shaded needles maximized foliar area to capture more light. Nitrogen concentrations were higher in shaded needles than in sun-exposed needles. This nitrogen distribution pattern was related to the high nitrogen cost of light interception and was assumed to improve light absorptance per needle mass of shaded needles. In contrast, in both 1- and 2-year-old needles, morphology was independent of prevailing light conditions; however, needle nitrogen concentrations were adjusted toward more effective light interception in 2-year-old foliage but not in 1-year-old foliage, indicating that acclimation of sun-adapted needles to shading takes more than one year. At the same time, needle aging was accompanied by accumulation of nonstructural carbohydrates (NSC), and increasing concentrations of needle carbon, suggesting a shift in the balance between photosynthesis and photosynthate export. The accumulation of NSC and carbon resulted in a dilution of the concentrations of other needle chemicals and explained the decline in needle nitrogen concentrations with increasing age. Thus, although morphological inadequacy to low light availabilities may partly be compensated for by modifications in needle chemistry, age-related changes in needle stoichiometric composition progressively lessen the potential for acclimation to low irradiance. A conceptual model, advanced to explain how environmental factors and age-related changes in the activities of needle xylem and phloem transport affect needle longevity, predicted that adaptation of needle morphology to irradiance during the primary growth period largely determines the fate of needles during subsequent tree growth and development.

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Nutrient contents and concentrations in relation to growth of Picea abies and Fagus sylvatica along a European transect.

Mineral nutrition of Norway spruce (Picea abies (L.) Karst.) and beech (Fagus sylvatica L.) was investigated along a transect extending from northern Sweden to central Italy. Nitrogen (N) concentrations of needles and leaves in stands growing on acid soils did not differ significantly between central Italy and southern Sweden (1.0 +/- 0.1 mmol N g(-1) for needles and 1.9 +/- 0.14 mmol N g(-1) for leaves). In both species, foliar N concentrations were highest in Germany (1.2 mmol N g(-1) for needles and 2.0 mmol N g(-1) for leaves) and decreased by 50% toward northern Sweden (0.5 mmol N g(-1)). Both species showed constant S/N and P/N ratios along the transect. Calcium, K and Mg concentrations generally reflected local soil conditions; however, Mg concentrations reached deficiency values in Germany. Leaf area per unit dry weight varied significantly along the transect with lowest values for Norway spruce recorded in northern Sweden and Italy (3.4 m(2) kg(-1)) and a maximum in central Europe (4.7 m(2) kg(-1)). A similar pattern was observed for beech. Despite the low variation in foliar N concentrations on the large geographic scale, local and regional variations in N concentrations equalled or exceeded the variation along the entire continental transect. Furthermore, nutrient contents (i.e., nutrient concentration x dry weight per needle or leaf) showed a greater variation than nutrient concentrations along the transect. Nitrogen contents of Norway spruce needles reached minimum values in northern Sweden (2.4 micro mol N needle(-1)) and maximum values in Denmark (5.0 micro mol N needle(-1)). The N content of beech leaves was highest in Denmark (242 micro mol N leaf(-1)). At the German site, foliar N content rather than N concentration reflected the seasonal dynamics of foliar growth and N storage of the two species. During foliage expansion, there was an initial rapid increase in N content and a decrease in N concentration. This pattern lasted for about 2 weeks after bud break and was followed by 6 weeks during which dry weight and N content of the foliage increased, resulting in a further decrease in N concentration. During summer, dry weight and N content of mature needles of Norway spruce increased further to reach a maximum in autumn, whereas N concentration remained constant. In spring, reallocation of N from 1- and 2-year-old needles was 1.5 and 1.0 micro mol N needle(-1), respectively. This remobilized N was a major source of N for the development of new needles, which had an N content of 1.5 micro mol N needle(-1) after bud break. The seasonal remobilization of N from old foliage decreased with increasing needle age. Needle N content and dry weight decreased progressively with age (1 micro mol N needle(-1) between age classes 2 and 5), whereas N concentrations remained constant. For Norway spruce, annual stemwood production was correlated with needle N content but not with foliar N concentration or with the total amount of N in the canopy. Interspecific and geographical differences in plant nutrition are discussed on the basis of competitive demands for C and N between growth of foliage and wood.

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Seasonal air and soil temperature effects on photosynthesis in red spruce (Picea rubens) saplings.

Net photosynthesis and stomatal conductance were measured in ten red spruce (Picea rubens Sarg.) saplings, growing near Ithaca, New York, throughout the early spring and late-fall growing periods. Gas exchange and daily minimum and maximum soil and air temperatures were also measured. Linear regression analysis showed that rates of net photosynthesis were positively correlated with both minimum daily soil and air temperatures but that minimum soil temperature was a better predictor of net photosynthesis. Moreover, net photosynthesis was more sensitive to changes in soil temperature than to changes in air temperature, and photosynthesis was approximately twice as sensitive to temperature changes during the fall than during the spring.

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Effects of cone-induction treatments on black spruce (Picea mariana) current-year needle development and gas exchange properties.

Both drought and root pruning (RP) increased the number of cones induced when black spruce (Picea mariana (Mill.) B.S.P.) grafts were injected with gibberellins A(4/7) (GA), but their effects on predawn shoot water potential and current-year needle development differed. Drought decreased predawn shoot water potential (Psi(pd)), but only during the period when irrigation was withheld, and it had no effect on the growth or gas exchange properties of current-year needles. Conversely, root pruning had little effect on Psi(pd), but it resulted in trees with smaller current-year needles that had lower nitrogen and chlorophyll concentrations and reduced rates of gas exchange up to the later stages of shoot elongation compared with needles of control trees. These findings are discussed in relation to potential effects on the development of induced cones in the following growth cycle.

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Seasonal photosynthetic responses to light and temperature in white spruce (Picea glauca) seedlings planted under an aspen (Populus tremuloides) canopy and in the open.

Photosynthetic light and temperature response curves were measured seasonally in seedlings of white spruce (Picea glauca (Moench.) Voss) grown for two years in the understory of aspen (Populus tremuloides Michx.) or in the open in central Alberta. Light-saturated rate of net photosynthesis, the optimum temperature for net photosynthesis, transpiration rate, photochemical efficiency, and stomatal and mesophyll conductances increased from spring to summer and declined thereafter, whereas dark respiration rate and compensation and saturation points were highest in spring. Depression of photosynthetic parameters was greater in open-grown seedlings than in understory seedlings during the periods in spring and autumn when night frosts were common. Net photosynthetic rates were similar in understory and open-grown seedlings in summer, but they were significantly lower in open-grown seedlings in spring and autumn. Significantly lower transpiration rates and stomatal conductances in open-grown seedlings than in understory seedlings were also observed at 15 and 25 degrees C in the autumn. Shoot and needle growth were less in open-grown seedlings than in understory seedlings. In summer, when irradiances were low in the aspen understory, understory white spruce seedlings maintained a positive carbon balance by decreasing their compensation and saturation points and increasing their photochemical efficiency compared to spring and autumn.

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Variability in leaf-level CO(2) and water fluxes in Pinus banksiana and Picea mariana in Saskatchewan.

We measured seasonal and canopy-level gas exchange in two stands of jack pine (Pinus banksiana Lamb.) and one stand of black spruce (Picea mariana (Mill.) B.S.P.) on relatively clear days from late May until mid-September 1994. Field measurements were made with a portable infrared gas analyzer, and laboratory measurements included photosynthetic oxygen evolution and needle chemical composition. Seasonally averaged light-saturated assimilation rates in the field were 4.0 micro mol m(-2) s(-1) in jack pine and 2.7 micro mol m(-2) s(-1) in black spruce. Rates of assimilation and transpiration were highest in midsummer. The seasonal pattern was especially pronounced for black spruce, probably because cold soil temperatures limited early season gas exchange rates in this species. Among stands, instantaneous water-use efficiency was highest in a young jack pine stand early in the season and higher in the upper canopy foliage than in the lower canopy foliage at all sites at the end of the season. Needles of young jack pine exhibited higher photosynthetic capacity, dark respiration and needle N concentrations than needles of trees at the old site. In both species, slight acclimation to shading was manifested by reductions in photosynthetic capacity in the lower canopy foliage. In both species, first-year needles had greater photosynthetic capacity than older needles but in situ rates of CO(2) assimilation in the field showed little difference among needle age classes. In both species, there was a strong correlation between assimilation and stomatal conductance, indicating that assimilation was highly stomatal limited and that environmental factors that alter conductance (e.g., VPD) have a strong influence on CO(2) and water fluxes, especially after early season thawing concludes.

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Organic and inorganic sulfur transport in the xylem sap and the sulfur budget of Picea abies trees.

Temporal changes in inorganic and organic sulfur compounds (sulfate, glutathione, cysteine, methionine) were analyzed in xylem sap of 40-year-old Norway spruce (Picea abies (L.) Karst.) trees growing on acidic soils at a healthy and a declining stand in the Fichtelgebirge (North Bavaria, Germany). Studies were carried out (1) to quantify glutathione (GSH) transport in the xylem of spruce, (2) to study the significance of reduced sulfur versus sulfate (SO(4) (2-)) transport in the xylem, and (3) to compare total sulfur (S) transport in the xylem with the amount of foliar uptake of SO(2) in an air-polluted environment. Glutathione was the main reduced S compound in the xylem ranging in concentration from 0.5 to 5 &mgr;mol l(-1). Concentrations of inorganic SO(4) (2-) in the xylem sap were up to 50 times higher than those of GSH ranging from 60 to 230 &mgr;mol l(-1). During the growing season, concentrations of all S compounds in the xylem were highest in May (up to 246 &mgr;mol l(-1)) and decreased during summer and fall (up to 21 &mgr;mol l(-1)). On average, SO(4) (2-) concentrations in xylem sap were 30% higher at the declining site compared with the healthy site. Diurnal changes in organic S compounds were significant for GSH and cysteine with high concentrations during the night and low concentrations during the day. Diurnal changes in inorganic concentrations were not significant. Xylem sap concentrations of SO(4) (2-) and cysteine were twice as high and GSH concentrations were tenfold higher in surface roots than in branches. At both sites, transport of organic S was low (up to 3% of total S) compared to transport of SO(4) (2-). Annual transport of total S in the xylem (SO(4) (2-) was the main component) ranged from 60 to 197 mmol tree(-1) year(-1) at the healthy site and from 123 to 239 mmol tree(-1) year(-1) at the declining site. Although gaseous uptake of SO(2) was estimated to be similar at both sites (38 mmol tree(-1) year(-1); Horn et al. 1989), the ratio between annual gaseous uptake of SO(2) and transport of S in the xylem was 1:4 and 1:5 at the healthy and declining sites, respectively.

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Effects of photoperiod and temperature on the timing of bud burst in Norway spruce (Picea abies).

We examined the effects of several photoperiod and temperature regimes imposed during the winter-spring period on the timing of bud burst in rooted cuttings of Norway spruce (Picea abies (L.) Karst.) grown in a greenhouse in Finland. The treatments were initiated in November and December after the cuttings had been exposed to natural chilling and freezing events. Irrespective of the treatments applied, time to bud burst decreased with increased duration of previous exposure to natural chilling and freezing events. Fluctuating day/night temperatures and continuous lengthening of the photoperiod hastened bud burst. Shortening the photoperiod delayed bud burst, suggesting that little or no ontogenetic development toward bud burst takes place during mild periods before the winter solstice. In the case of climatic warming, this phenomenon may prevent the premature onset of growth that has been predicted by computer simulations with models that only consider temperature regulation of bud burst.

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Flower induction treatments have no effects on seed traits and transmission of alleles in Picea glauca.

Flower induction methods-hormone application or exposure to physiological stress, or both-are used routinely for shortening breeding cycles and increasing seed production in white spruce (Picea glauca (Moench) Voss). The objectives of this study were: (1) to evaluate the effects of flower induction on seed yield and quality in white spruce; and (2) to determine if flower induction treatments affect the maternal contribution to offspring. We assessed the effects of flower induction treatments, which consisted of gibberellin A(4/7) (GA(4/7)) and naphthaleneacetic acid (NAA) stem injections, on allele segregation for 28 clones, number of seeds per cone, number of sound seeds per cone, seed weight, and the germination rate of a subset of clones. Flower induction treatments did not affect any of the phenotypic traits examined. No increase in segregation distortion in allozyme loci following flower induction treatments was observed.

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Dynamics of change in stomatal response and water status of Picea abies during a persistent drought period: a contribution to the traditional view of plant water relations.

Four experiments on the simulation of a persistent drought period were carried out with cloned Picea abies (L.) Karst. trees: two in the field under varying weather conditions and two in a climate chamber under variously manipulated humidity conditions. Patterns of diurnal dynamics in gas exchange rates and water potential were monitored and analyzed. The first phase of the drought was characterized by relatively high daily maxima for photosynthesis and transpiration. With decreasing humidity during the day, the values dropped steeply, and the declines were larger and occurred earlier on each passing day of the drought period. When soil water potential was lower than -2000 hPa, maximum stomatal aperture was greatly reduced despite a humid atmosphere. Under these conditions, rates of photosynthesis and transpiration decreased less steeply from the daily maxima and differences between maxima and minima were small. In the field, the daily sums of transpiration and photosynthesis were more dependent on atmospheric conditions than on soil water potential. In the growth chamber experiments, the daily sums of transpiration and photosynthesis decreased continuously as the soil dried, at first steeply until a soil water potential of -2000 hPa was reached, then slowly. Predawn water potential values fluctuated under field conditions, but tended to decrease with time, whereas needle osmotic potential increased slightly. Because relative humidities did not reach 100% in the growth chamber, predawn water potentials of plants in the growth chamber were never higher than -1.0 MPa although the soil was saturated. In the experiment with a high average air humidity during the daily stress period, relatively high predawn water potentials were maintained until lower soil water potentials of -8000 hPa were reached. Results were used to assess the importance of evaporative demand versus soil drying on stomatal responses within the context of current concepts of plant water relations. The observed trends in diurnal dynamics can be explained solely by the interdependency of leaf conductance and water potential. Stomata react directly to the ratio of water supply to demand. The central role of peristomatal transpiration in this system is emphasized.

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Effect of season and soil treatments on carbohydrate concentrations in Norway spruce (Picea abies) mycorrhizae.

We studied effects of season and soil treatments (watering, acidification, liming and combinations of these treatments) on soluble carbohydrates of mycorrhizal roots of Norway spruce (Picea abies (L.) Karst.). Arabinose, arabitol, fructose, glucose, inositol, lactose, mannitol, pinite, quinate, raffinose, shikimate, stachyose and trehalose were identified by HPLC. Concentrations of inositol, lactose and pinite were constant throughout the year, whereas concentrations of raffinose, stachyose and trehalose were higher in winter than in summer, and concentrations of glucose, fructose and mannitol increased from February to September. Soil acidification and liming had no effect on the annual mean concentrations of fructose, glucose, lactose, pinite, raffinose and stachyose. Liming increased quinate concentrations and decreased arabitol concentrations. Annual mean concentrations of arabinose and mannitol decreased in response to soil acidification. Annual mean concentrations of inositol increased in response to irrigation. None of the soil treatments affected the mean annual concentration of trehalose.

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Freezing cycles enhance winter injury in Picea rubens.

We examined changes in chlorophyll absorbency in red spruce (Picea rubens Sarg.) foliage in response to simulated freezing cycles. Current-year branch tips were collected from 16 trees on January 8, January 20, February 8 and February 26, 1996. Tissue was subjected to freezing cycle treatments with a minimum of -35 degrees C and a maximum of 3 degrees C for a one-cycle treatment, and -9, -6, -3, 0 or 3 degrees C for four-cycle treatments. Samples were frozen at a rate of 5 degrees C h(-1), and warmed at 12 to 15 degrees C h(-1). Controls were held at -9 degrees C. Temperatures during the three-day periods preceding each sample date averaged -18, 4.7, -9.6 and 3.7 degrees C, respectively. On January 8, treated trees showed no significant (P > 0.1) increase in the breakdown of chlorophyll, as measured by the ratio of chlorophyll a absorbency (435 nm) to phaeophytin a absorbency (415 nm), compared with control branch tips. On later sampling dates, seven trees consistently exhibited needle reddening and nine exhibited few symptoms (< 10% of total needle surface reddened) after four-cycle treatments. On February 26, chlorophyll degradation in trees with needle reddening differed (P < 0.05) from the control by 26, 26, 16, 14 and 15% for the 3, 0, -3, -6 and -9 degrees C maxima, respectively. No detectable chlorophyll degradation occurred after a one-cycle treatment in any trees on any sampling date. Freezing cycles with sub-zero maxima and a -35 degrees C minimum enhanced winter injury in red spruce after a midwinter thaw had rendered the trees susceptible to freezing damage.

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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.

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