Diurnal variation in xylem hydraulic conductivity in white ash (Fraxinus americana L.), red maple (Acer rubrum L.) and red spruce (Picea rubens Sarg.)

Xylem hydraulic conductivity and percentage loss of conductivity (PLC) were measured on a ring-porous ( Fraxinus americana L., white ash), a diffuse porous ( Acer rubrum L., red maple) and a coniferous ( Picea rubens Sarg., red spruce) tree species in a temperate deciduous forest in central Massachusetts, USA. Measurements were made on current and 1-year-old branch segments in the afternoon and on the following morning. Afternoon PLC was 45 to 70% for the current year's extension growth in both white ash and red maple. Morning PLC was significantly lower (10–40%). Conductivity also varied diurnally suggesting, on average, a 50% recovery from cavitation overnight. Red spruce showed lower PLC and conductivity and a less pronounced night-time recovery. Diurnal variation in hydraulic conductivity and PLC suggests that embolism removal occurred in all three species despite the existence of tension within the xylem. Further evidence for embolism removal was observed with an in situ double-staining experiment in which dyes were fed to a transpiring branch during the late afternoon and the following morning. Examination of stem cross-sections showed that a larger number of vessels were conductive in the morning than on the preceding afternoon. Results of this study suggest that hydraulic capacity is highly dynamic and that conductivity measurements reflect a balance between two processes: cavitation and embolism removal.     

Biogeochemical controls on aluminum chemistry in the O horizon of a red spruce (Picea rubens Sarg.) stand in central Maine,USA

This study examined the biotic and abiotic processes controlling solution chemistry and cycling of aluminum (Al) in the organic horizons of a northern coniferous forest ecosystem. A mass balance budget indicated that aboveground inputs of Al to the O horizon averaged 0.9 kg ha^–1 1 yr^–1, with major inputs accounted for by litterfall (69%), followed by precipitation (21%), and net canopy throughfall plus stemflow (10%). Estimated leaching losses of Al from the O horizon averaged 2.1 kg Al ha^-1 yr1. We hypothesize that the difference between measured Al inputs and outputs can be accounted for by Al release from weathering of soil minerals admixed into the O horizon. Variations in O horizon solution Al chemistry were influenced by a number of factors, including pH, Al equilibria with different solid-phase organic exchange sites, and Al complexation with humic ligands in soil solution.     

Potential for red spruce (Picea rubens Sarg.) establishment from natural seed dispersal in old fields adjacent to forest stands

Red spruce can establish on abandoned agricultural land only from seed, unlike in logged environments where red spruce forests can establish from seedlings that survived the harvest. In this study we collected seeds from the seed rain and from soil cores to examine the abundance and distribution of the red spruce seed rain and the seed bank along a gradient from the forest interior to adjacent old fields in Greenwich, Prince Edward Island National Park (PEINP), Canada. In addition, we examined the temporal abundance and distribution of the seed rain through the seed-fall season. We subsequently tested the germination potential of red spruce seeds from the forest portion of our sites. Because red spruce seeds are usually viable for only one year, we expected the seed bank to be either very small or non-existent. We found 39 red spruce seeds in the seed bank, distributed over 13.1% of the sampled area; 37 of these seeds were definitely nonviable. From the seed rain, we collected 224 red spruce seeds, distributed over 29.4% of the sampled area; 213 of these seeds were definitely nonviable. Nearly all of the red spruce seed rain fell within the forest; differences in abundance among collection dates were not statistically significant. Our seeds had a low germination rate of 0.004%, in part because red and black spruce commonly hybridise in our study area. Despite the low rates of red spruce seed dispersal and viability, the presence of red spruce saplings in our two longest-abandoned fields shows that the species is able to slowly colonise some old fields in PEINP.     

Carbon dioxide assimilation and growth of red spruce (Picea rubens Sarg.) seedlings in response to ozone,precipitation chemistry,and soil type

Summary The influence of ozone, mist chemistry, rain chemistry, and soil type on CO₂ assimilation and growth of red spruce (Picea rubens Sarg.) seedlings was investigated over a 4-month period under controlled laboratory and glasshouse conditions. Growth was evaluated through interval estimates of aboveground relative growth rates (RGR) and the partitioning of biomass components at harvest to root, stem, and needle fractions. Precipitation chemistry treatments and O₃ exposure dynamics were based on reported characteristics of air chemistry and/or deposition in high-elevation forests of eastern North America. The two soils were collected from Camels Hump in the Green Mountains of Vermont and Acadia National Park on the Maine coast. Soil from Acadia had higher organic content, higher levels of extractable base cations, and lower levels of extractable aluminum and heavy metals. The only treatment variables that consistently influenced the growth of P. rubens were soil type and rain chemistry. In comparison with seedlings grown in soil from Acadia National Park, those grown in Camels Hump soil had significantly less needle (27%), stem (33%), and root (26%) biomass at harvest and statistically lower aboveground RGR within 2 months after initiation of the treatments. Seedlings grown in Camels Hump soil had significantly higher levels of aluminum (6.5X), copper (1.4X), and nickel (2.7X) in new needle tissue. The only influence of precipitation chemistry on the growth of P. rubens was a pattern of greater root and shoot biomass in seedlings experiencing the more acidic rain treatments. Interactive effects among the main treatment variables (e.g., acidic mist and O₃, acidic rain and soil type) on seedling growth were not notable. Rates of CO₂ assimilation and transpiration on a per gram needle dry weight basis [mol·g^-1·s^-1] were not influenced by any of the main treatment variables or their interaction. Because neither soil type nor precipitation chemistry influenced the efficiency of CO₂ assimilation per gram dry weight of needle tissue, the physiological mechanism underlying the growth response of P. rubens is attributed to a change in either whole-plant allocation of carbon resources or a direct toxic effect in the rhizosphere on root growth.     

Temporal patterns of inorganic nitrogen uptake by mature sugar maple (Acer saccharum Marsh.) and red spruce (Picea rubens Sarg.) trees using two common approaches

Background: Plant uptake of nitrogen influences many ecosystem processes, yet uptake by trees in northern forests of the United States has not been quantified throughout the growing season.

Aims: To measure NH₄ ⁺ and NO₃ ^? uptake by mature sugar maple (Acer saccharum) and red spruce (Picea rubens) trees during the early, mid and late growing season.

Methods: At Hubbard Brook Experimental Forest, New Hampshire, we used two approaches to measure nitrogen uptake capacity by mature trees: an in situ depletion method using intact roots and an ex situ ¹⁵N tracer method using excised roots.

Results: NH₄ ⁺ uptake was greater than NO₃ ^? for both methods and tree species (P < 0.05). NH₄ ⁺ uptake was lowest during the early growing season, while NO₃ ^? uptake was lowest during the late growing season. Measured rates of NH₄ ⁺ uptake were 2–3 orders of magnitude greater using the in situ depletion method compared with the ex situ ¹⁵N tracer method.

Conclusions: These results demonstrate seasonal differences in nitrogen uptake by two dominant tree species in a northern forest and show that the method employed can significantly impact measured rates of uptake, which could have implications for understanding the magnitude of plant nitrogen uptake and for cross-study comparisons of this process.     

Carbohydrate requirements for the development of black spruce (Picea mariana (Mill.) B.S.P.) and red spruce (P. rubens Sarg.) somatic embryos

Different carbohydrates were investigated for somatic embryo development of black spruce and red spruce. They were tested in a basal maturation medium consisting of Litvay's salts at half-strength containing 1 g l^-1 glutamine, 1 g l^-1 casein hydrolysate, 7.5 M abscisic acid, and 0.9% Difco Bacto-agar. A comparison of different sucrose concentrations showed that 6% was optimal for embryo development. Among the nine carbohydrates tested, sucrose, fructose, glucose, maltose, and cellobiose supported embryo development while arabinose, mannitol, myo-inositol, and sorbitol did not. A comparison of sucrose, glucose, and fructose at three concentrations showed that the general pattern of response for both species followed concentration expressed as a percentage, independent of the molarity of carbohydrate in the medium. Interspecific differences were observed concerning carbohydrate requirements. For red spruce, 6% fructose was found best for embryo development, while no such preference was observed for black spruce. No significant difference was observed in the number of embryos produced with 6% sucrose or 3% sucrose plus an equimolar concentration of either mannitol, sorbitol, or myo-inositol in the maturation medium, suggesting that the effect of the carbohydrate on the maturation was partly osmotic.     

Polyamines and their biosynthetic enzymes during somatic embryo development in red spruce (<Emphasis Type="Italic">Picea rubens</Emphasis> Sarg.)

Summary The major objective of this study was to determine if the observed changes in polyamines and their biosynthetic enzymes during somatic embryo development were specifically related to either the stage of the embryo development or to the duration of time spent on the maturation medium. Somatic embryos of red spruce (Picea rubens) at different developmental stages, grown in the embryo development and maturation media for various lengths of time, were separated from the associated subtending tissue (embryogenic and the suspensor cell masses) and analyzed for their polyamine content as well as for polyamine biosynthetic enzyme activities. Polyamine content was also analyzed in embryos representing different stages of developmentthat were collected from the sam culture plate at the same time and the subtending tissue surrouding them. Putrescine was the predominant polyamine in the pro-embryogenic tissue, while spermidine was predominant during embryo development. Significant changes in spermidine/putrescine and spermine/putrescine ratios were observed at all stages of embryo development as compared to the pro-embryogenic cell mass. Changes in the ratios of various polyamines were clearly correlated with the developmental stage of the embryo rather than the period of growth in the maturation medium. Whereas the activities of both ornithine decarboxylase and arginine decarboxylase increased by week 3 or 4 and stayed high during the subsequent 6 wk of growth, the activity of S-adenosylmethionine decarboxylase steadily declined during embryo development.     

Blue intensity as a temperature proxy in the eastern United States: A pilot study from a southern disjunct population of Picea rubens (Sarg.)

Annual surface air temperatures across the eastern United States (US) have increased by more than 1 °C within the last century, with the recent decades marked by an unprecedented warming trend. Tree-rings have long been used as a proxy for climate reconstruction, but few truly temperature-sensitive trees have been documented for the eastern US, much less the Appalachian Mountains in the Southeast. Here, we measure blue intensity (BI) and ring width (RWI) in red spruce growing at the southernmost latitudinal range margin of the species on the North Carolina-Tennessee border to test the efficacy of using either metric as a temperature proxy in the eastern US. The BI and RWI chronologies spanned 1883–2008 and had an interseries correlations of 0.42 and 0.54, respectively, but time series were trimmed to the period 1950–2008 due to low sample depth. We discovered strong, positive, and stable correlations between both current-year early fall (September–October) T_max (r = 0.62; p < 0.001) and T_mean (r = 0.51; p < 0.001) and ΔBI during the period 1950–2008, but found no significant relationships between temperature and RWI. We show BI metrics measured in red spruce to be a promising temperature proxy for the southern Appalachian Mountain region. Future research should focus on testing [1] the efficacy of using BI on red spruce collected from across the species range, and [2] the potential for using BI as a temperature proxy in other conifers distributed in the eastern US.     

Growth of adult Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica L.) under free-air ozone fumigation

This study attempted to detect the impact of ozone on adult trees of Norway spruce ( Picea abies [L.] Karst.) and European beech ( Fagus sylvatica L.) in an experimental mixed stand in Southern Bavaria, Germany. The aim was to examine whether there is a decrease in growth when trees are exposed to higher than atmospheric concentrations of ozone. This exposure was put into effect using a free-air fumigation system at tree crown level. Growth analysis was carried out on a group of 47 spruce and 36 beech trees, where radial stem increment at breast height - a sensitive index for stress - was measured. The ozone monitoring system allowed values to be obtained for the accumulated ozone exposure (SUM00) of each individual tree, so that their radial increment over three years could be correlated with the corresponding ozone exposure for the same time period. Correlation and regression analysis were then carried out to test the influence of ozone on diameter increment. In both spruce and beech, the initial stem diameter was the most influential factor on radial increment in the following year. A linear model was applied, including the diameter of the preceding year and the ozone exposure of the current year as predicting factors. For spruce trees, a significant negative influence of ozone exposure was found. In contrast, no significant ozone effect on diameter increment of beech was detected. The effect of ozone stress on a large spruce tree can lead to a decrease in potential radial increment of 22 %. The results are discussed in relation to other stress factors such as drought and lack of light.     

Cytochrome and alternative pathway respiration in white spruce (Picea glauca) roots. Effects of growth and measurement temperature

Interactions between growth temperature and measurement temperature were examined for their effects on white spruce [ Picea glauca (Moench) Voss] root respiration. Total dark respiration rates increased with measurement temperature and were unaffected by growth temperature. Partitioning of respiratory electron flow between the cytochrome and alternative pathways was also unaffected by growth temperature. The proportion of respiration mediated by the alternative pathway was constant at measurement temperatures between 4°C and 18°C, but was increased at higher temperatures. Changes in alternative pathway activity were paralleled by changes in capacity, and the alternative pathway was almost fully engaged at all temperatures. Roots grown at low temperature displayed higher carbohydrate levels than roots grown at higher temperatures, but respiration rate was unaffected. Spruce root respiration did not appear to acclimate to growth temperature, and the alternative pathway was not preferentially engaged at low temperature.     

The effect of open-air fumigation with SO2 and O3 on carbohydrate metabolism in Scots pine (Pinus sylvestris) and Norway spruce (Picea abies)

The carbohydrate metabolism of the needles of Scots pine (Pinus sylvestris) and Norway spruce (Picea abies) has been examined in trees that were exposed to SO₂, and O₃, in an open-air fumigation experiment located in the Liphook forest in southern England. Two-year-old seedlings were planted in 1985 in seven experimental plots. Five plots received fumigation treatments of SO₂, O₃ or a combination of these gases to give a 2 × 3 factorial design with one additional ambient plot Fumigation with SO₂, occurred from May 1987 to December 1990 and O₃, fumigation occurred from March to December 1988, May to December 1989 and February to December 1990. Five samples of needles for investigation of carbohydrate metabolism were taken between February and July 1989. The concentrations of soluble carbohydrates (including sucrose and hexoses) were greatly reduced in the needles taken from Scots pine growing in the treated plots, and were also reduced, but to a lesser extent, in the needles taken from Norway spruce. Little variation in the concentration of starch in the needles of either species was detected. The activities of the two final enzymes of sucrose synthesis, sucrose phosphate synthase and sucrose 6-phos-phate phosphatase, were greatly reduced in the needles of Scots pine and were also reduced, but to a lesser extent, in the needles of Norway spruce in the fumigated plots. These reductions could be correlated with decreases in rates of photosynthetic CO₂ assimilation determined by independent groups of researchers working on the Liphook site.     

The effect of long term CO2 enrichment on the growth,biomass partitioning and mineral nutrition of Sitka spruce (Picea sitchensis (Bong.) Carr.)

Sitka spruce [Picea sitchensis (Bong.) Carr.] seedlings were grown for 3 years in an outside control plot or in ambient (355 mol mol^-1) or elevated (ambient + 350 mol mol^-1) atmospheric CO₂ environments, within open top chambers (OTCs) at the Institute of Terrestrial Ecology, Edinburgh. Sequential harvests were carried out at the end of each growing season and throughout the 1991 growing season, five in all. Plants grown in elevated CO₂ had, (i) 35 and 10% larger root/shoot ratios at the end of the first and third season, respectively, (ii) significantly higher summer leader extension relative growth rates, which declined more rapidly in early autumn than ambient grown plants, (iii) after three growing seasons a significantly increased mean annual relative growth rate, (iv) consistently lower foliar nutrient concentrations, and (v) after two growing seasons smaller total projected needle areas. Plants grown inside OTCs were taller, heavier and had a smaller root/shoot ratio than those grown outside the chambers. There was no effect of CO₂ concentration on Sitka spruce leaf characteristics, although leaf area ratio, specific leaf area and leaf weight ratio all fell throughout the course of the 3 year experiment.     

Effects of varied soil nitrogen supply on Norway spruce (Picea abies [L.] Karst.)

During a seven-month period the effect of different nitrogen (N) availability in soil on growth and nutrient uptake was studied in three-year-old Norway spruce (Picea abies [L.] Karst.) trees. The plants were grown in pots on N-poor forest soil supplied with various amounts and forms (inorganic and organic) of N. Increasing supply of inorganic N (as NH₄NO₃) increased the formation of new shoots and shoot dry weight. The root/shoot dry weight ratio of new growth was drastically decreased from 1.6 in plants without N supply to 0.5 in plants supplied with high levels of NH₄NO₃. This decrease in root/shoot dry weight ratio was associated with distinct changes in root morphology in favour of shorter and thicker roots. The addition of keratin as organic N source did neither affect growth nor root morphology of the trees. The amount of N taken up by plants was closely related to the supply of inorganic N, and trees supplied with highest levels of NH₄NO₃ also had the highest N contents in the dry matter of needles and roots. In contrast, N contents in needles of trees grown without additional N, or with keratin supply, were in the deficiency range. Supply of NH₄NO₃ decreased the contents of phosphate (P) and potassium (K) and therefore markedly increased N/P and N/K ratios in the needles. On the other hand, the contents of calcium (Ca), magnesium (Mg), and manganese (Mn) in the needles were increased in the plants supplied with inorganic N, suggesting high soil availability and promotion of uptake of these divalent cations by high nitrate uptake. The observed effects on root/shoot dry weight ratio, root morphology, and mineral nutrient composition of the needles indicated that high inorganic N supply may increase above-ground productivity but at the same time decrease the tolerance of trees against soil-borne (e.g. deficiency of other mineral nutrients) stress factors. Deceased 21 September 1996 Deceased 21 September 1996     

Photosynthetic performance, chloroplast pigments and mineral content of Norway spruce (Picea abies (L.) Karst.) exposed to SO2 and O3 in an open-air fumigation experiment

Photosynthetic performance, mineral content and chloroplast pigments were investigated in August-September 1988 and 1989 in Norway spruce trees (Picea abies (L.) Karst.) exposed to SO₂, and O₃ in an open-air fumigation facility at Liphook, England. The data do not suggest a treatment effect on the mineral content of the needles in terms of nutrient leaching from the foliage. In addition, there were no direct SO₂ and/or O₃ effects on the content and/or composition of the chloroplast pigments. However, the long-term application of SO₂ resulted in a depression of net photosynthesis under light saturation and ambient CO₂ (A ₃₄₀) which was probably caused by a treatment-related depression of the carboxylation efficiency (CE). In 1989, the supposed treatment effects were apparently masked by an insufficient N-supply and probably also by low water availability during summer. However, fumigation appeared to accelerate an N-deficiency-related decrease of CE, stomatal closure and the age-dependent development of the chlorophyll content of the needles. In 1989, an observed depression of the photosynthetic capacity (A₂₅₀₀) was in part accompanied by a decrease in light use efficiency (α), suggesting an enhanced photosensitivity resulting from the impact of several possible interacting stresses (drought, N deficiency and fumigation). The results support the general conclusion that long-term low-level SO₂ dosage adversely affects the photosynthetic performance of the needle, whether directly or indirectly, and may also interact with other environmental stresses. The findings of our investigations are discussed with regard to the hypothesis of forest decline in the mountain regions of the Fichtelgebirge (north-eastern Bavaria, Germany).     

An attempt to predict long-term effects of atmospheric nitrogen deposition on the yield of Norway spruce stands (Picea abies (L.) Karst.) in southwestern Sweden

Long-term field experiments in Norway spruce stands on fertile sites (site indices 27–35 m) in southwestern Sweden were analysed with respect to volume increment. Three treatments were included (0=No fertilization, N = Fertilization with N, NP = Fertilization with N and P).Volume growth was monitored for 18 years in 10 blocks. No significant differences in annual volume increment between the treatments were detected. Volume increments in the N treatment were 97%, 99% and 107% as high as those in the 0 treatment for the periods 1–5, 6–10 and 11–15 years after the first fertilization. Corresponding values for the NP treatment were 104%, 108% and 110%, indicating that P has a small positive effect.The amount of N-fertilization would correspond to an annual N deposition of 20 kg ha^-1 during the next 30 years in southwestern Sweden. For this period, the results imply that this N deposition would not affect the growth of Norway spruce stands on fertile sites.