Nutrient concentrations in roots, leaves and wood of seedling and mature sugar maple and American beech at two contrasting sites

Differences in sensitivity to soil conditions across tree species and developmental stage are important to predicting forest response to environmental change. This study was conducted to compare elemental concentrations in leaves, stems, and roots of (1) sugar maple (Acer saccharum Marsh.) seedlings vs. mature trees and (2) mature sugar maple vs. mature American beech (Fagus grandifolia Ehrh.) in two sites that differ in soil base saturation and pH. Both sites are located in Huntington Forest, NY, USA; one site (hereafter ‘H’) has higher soil pH and Ca, Mg, and Mn concentrations than the other site (hereafter ‘L’). Sugar maple growth at H (14.8 cm² year⁻¹ per tree) was much greater than at L (8.6 cm² year⁻¹ per tree), but the growth of beech was not different between the two sites. Leaves, roots, and stem wood of mature beech trees and sugar maple seedlings and mature trees were sampled for nutrient analysis. Foliar Ca, K, and Al concentrations were positively correlated with soil elements, but Mn concentrations were negatively correlated. Sugar maple differed more than beech between sites in foliar K and Mn concentrations. Root Mg and P concentrations reflected soil chemistry differences, in contrast to foliar concentrations of Mg and P, which were indistinguishable between the sites. In sugar maple, seedlings differed more than in mature trees in nutrient concentrations in roots, especially for Mg and Mn. Although beech was not as responsive to nutrient availability as sugar maple in foliar and root nutrient concentrations, Ca and Mg concentrations in beech wood were higher in H (52% higher for Ca and 68% higher for Mg), while sugar maple did not differ between sites. Sugar maple regeneration failure on acidic soils in the same region is consistent with our finding that sugar maple seedlings were very sensitive to nutrient availability. This sensitivity could ultimately contribute to the replacement of sugar maple by American beech in regions of low pH and base cations if base cation leaching by anthropogenic deposition and tree harvesting continues.     

Effects of enhanced nitrogen deposition on foliar chemistry and physiological processes of forest trees at the Bear Brook Watershed in Maine

Effects of enhanced nitrogen deposition on nutrient foliar concentrations and net photosynthesis of sugar maple (Acer saccharum Marsh.), American beech (Fagus grandifolia Ehrh) and red spruce (Picea rubens Sarg.) were evaluated at the Bear Brook Watershed in Maine (BBWM). The BBWM is a paired-watershed forest ecosystem study with one watershed treated since 1989 with bimonthly dry ammonium sulfate ((NH₄)₂SO₄) additions at a rate of 25.2 kg N ha⁻¹ year⁻¹, while the other watershed serves as a reference. The (NH₄)₂SO₄ treatment resulted in significant increases in foliar N concentrations for all three species and significant reductions in foliar Ca, Mg and Zn concentrations for American beech and red spruce. Treatment effects on foliar concentrations of other nutrients were not significant in any species. Despite higher N concentrations in all species, only treated sugar maple showed significantly higher photosynthetic rates. The non-response in net photosynthesis to higher foliar N in American beech and red spruce might be attributed to their low foliar Ca and/or Mg concentrations. Higher net photosynthetic rates in sugar maple might be explained by the higher foliar N and by the ability of this species to maintain an adequate Ca and Mg supply. Results suggested that nutrient imbalances due to inadequate supply of Ca and Mg might have counteracted a potential increase in net photosynthesis induced by higher N concentrations in American beech and red spruce.     

The Effect of Landfill Leachate Spraying on Foliar Nutrient Concentrations and Leaf Transpiration in a Northern Hardwood Forest, Canada

The effect of municipal solid waste (MSW) leachate spray irrigationon a mature northern hardwood forest was investigated. Canopyfoliar samples and stem increment cores were collected fromtwo indicative species, sugar maple (Acer saccharum Marsh.)and American beech (Fagus grandifolia Ehrh.), within each ofa heavily sprayed, lightly sprayed and control area. Foliarconcentrations of N and P were significantly higher in bothmaple and beech foliage within the sprayed areas when comparedto an unsprayed area (control). Levels of Mg and K were markedlyhigher in maple but not beech foliar samples within the heavilysprayed areas when compared to foliage sampled within the unsprayedcontrol. While no significant trends were observed within themaple foliage, both Fe and B levels increased significantlyin beech foliar samples obtained from within the heavily sprayedarea in comparison to foliage samples from the control. Directporometric measurements of the transpiration rate and diffusive(stomatal) resistance of canopy and understory plant leavesrevealed a significant increase in diffusive resistance anda decline in transpiration rate with leachate spraying. Afterfour years of spraying a significant effect of leachate applicationon radial stem growth of both maple and beech trees has notbeen observed.     

Sugar maple (Acer saccharum Marsh.) decline during 1979–1989 in northern Pennsylvania

Sugar maple decline has been observed in northern Pennsylvania since the early 1980s. We investigated the interactions between soil moisture stresses in sugar maple and other factors, such as soil chemistry, insect defoliation, geology, aspect, slope, topography, and atmospheric deposition. In the summer of 1998, we sampled 28 sugar maple (Acer saccharum Marsh.) plots drawn from the USFS Forest Inventory and Analysis (FIA) plots, containing declining and non-declining sugar maple trees across northern Pennsylvania for a variety of soil physical and chemical parameters, site characteristics, and tree health. Foliage from declining plots was found to have significantly lower base cations and higher Mn as compared to that from non-declining plots. Soils in declining plots had lower base cations and pH, a Ca:Al⁻≤1, lower percent clay and higher percent sand and rock fragments than soils on non-declining plots, suggesting that trees on declining plots are at risk of nutritional and drought stress. Regression relationships between foliar and soil chemistry indicated that foliar nutrition was highly correlated with soil chemistry in the upper 50 cm of the soil. Declining sugar maple plots in this study occurred at higher elevations on sandstone dominated geologies. Soils were found to be base poor-sandy soils that contained high percentages of rock fragments. Soils below 50 cm on declining plots had lower soil pH and foliar chemistry indicated lower foliar base cations. A trend, while not significant was found with declining plots experiencing a greater number of and more severe insect defoliations.     

Spatial coexistence of American beech and sugar maple regeneration in post-harvest northern hardwood forests

Context

Fine scale regeneration patterns of coexistent species are influenced by regeneration mechanisms and microsite requirements. Spatial patterns may be either disjunct or overlapping, which will determine competitive effects and microsite dominance, and future forest composition.

Aims

Using American beech (Fagus grandifolia Ehrh.) and sugar maple (Acer saccharum Marshall) as an example, three hypotheses were tested: (1) random beech spatial patterns, (2) clumped spatial patterns of small sugar maple seedlings, and (3) disjunct beech and sugar maple patterns.

Methods

Individual stems were sampled in a contiguous grid of 1-m² quadrats across a 576-m² area at three sites. Densities were separated into three height classes (≤30 cm, 30–90 cm, and?>?90 cm, ≤4 cm diameter at breast height). Spatial statistics and regression were used to analyze spatial patterns and correlations.

Results

Beech and seedling sugar maple patterns were patchy, rejecting the first and not rejecting the second hypotheses. Hypothesis three was rejected because patches of the two species overlapped with advance regeneration beech overtopping sugar maple.

Conclusion

Patchy patterns of advance regeneration beech and post-harvest sugar maple establishment suggest spatiotemporal niche partitioning. Beech had a competitive height advantage following harvest, but sugar maple still occurred in beech-free patches and beneath overtopping beech at a fine scale. Self-replacing beech patterns will ensure the species will continue dominance unless a selective chemical or manual treatment is applied that removes beech and releases sugar maple.     

Chlorophyll content monitoring in sugar maple (Acer saccharum)

We conducted two experiments to determine the usefulness of a chlorophyll content meter (CCM) for the measurement of foliar chlorophyll concentration in sugar maple (Acer saccharum Marsh.) in the fall color period. In Experiment 1, four sugar maple trees were visually assigned to each of four fall foliage color categories in October 1998. On four dates in the fall of 1999, leaves were taken from the trees and analyzed for chlorophyll concentration by absorbance of pigment extracts and by determination of the chlorophyll content index (CCI) with a CCM. The two measures of chlorophyll concentration were strongly correlated (P < 0.001, r2 = 0.72). In Experiment 2, the CCI of leaves from sugar maple trees subjected to one of four fertilization treatments (lime, lime + manure, lime + 10:10:10 N,P,K fertilizer and an untreated control) were determined with a CCM. Treatment effects were distinguishable between all pairwise comparisons (P < 0.001), except for the lime versus lime + NPK fertilizer treatments.     

Effects of chronically elevated nitrogen and sulfur deposition on sugar maple saplings: Nutrition,growth and physiology

At the Bear Brook Watershed in Maine (BBWM), we examined the effects of long-term experimentally elevated N and S deposition on foliar chemistry, growth, and photosynthetic capacity of sugar maple (Acer saccharum) saplings. The BBWM is a paired watershed system; one watershed has been acidified bimonthly with granular ammonium sulfate ((NH₄)₂SO₄) since 1989. The adjacent watershed is used as a reference. We observed a 56% increase in foliar Al and a 25% reduction in foliar Ca for sugar maple saplings on the treated watershed compared to reference. Foliar N (+15%), P (+10%), and K (+15%) were significantly elevated in treated saplings. Along with changes in foliar nutrients, there were significant differences in photosynthetic capacity.     

Stem girdling manipulates leaf sugar concentrations and anthocyanin expression in sugar maple trees during autumn

To better understand the effects of sugar accumulation on red color development of foliage during autumn, we compared carbohydrate concentration, anthocyanin expression and xylem pressure potential of foliage on girdled versus non-girled (control) branches of 12 mature, open-grown sugar maple (Acer saccharum Marsh.) trees. Half of the study trees were known to exhibit mostly yellow foliar coloration and half historically displayed red coloration. Leaves from both girdled and control branches were harvested at peak color expression (i.e., little or no chlorophyll present). Disruption of phloem export by girdling increased foliar sucrose, glucose and fructose concentrations regardless of historical tree color patterns. Branch girdling also increased foliar anthocyanin expression from 50.4 to 66.7% in historically red trees and from 11.7 to 54.2% in historically yellow trees, the latter representing about a fivefold increase compared with control branches. Correlation analyses indicated a strong and consistent relationship between foliar red coloration and sugar concentrations, particularly glucose and fructose, in both girdled and control branches. Measures of xylem pressure potentials confirmed that girdling was a phloem-specific treatment and had no effect on water transport to distal leaves. Results indicate that stem girdling increased foliar sugar concentrations and enhanced anthocyanin expression during autumn in sugar maple foliage. Native environmental stresses (e.g., low autumn temperatures) that reduce phloem transport may promote similar physiological outcomes.     

Leaf nutrition and photosynthetic performance of sugar maple (Acer saccharum) in stands with contrasting health conditions

Leaf nutrition and photosynthetic performance of sugar maple (Acer saccharum Marsh.) were compared between two sugar maple stands in northwestern Vermont with contrasting health conditions as indicated by annual basal area growth, degree of crown dieback, and foliar appearance. Observations made during the diurnal cycle of both stands showed no apparent leaf water stress. In both stands, leaves had similar concentrations of major non-structural carbohydrates (starch and sucrose). Over two consecutive growing seasons (1991 and 1992), we consistently observed lower leaf Ca and Mg concentrations in the declining stand than in the healthy stand. Compared with the healthy stand, lower leaf chlorophyll concentrations and apparent leaf chlorosis were observed in the declining stand, and some trees had very low foliar Ca and Mg concentrations (0.31 +/- 0.03% and 0.09 +/- 0.01%, respectively). Trees in the declining stand had lower light-saturated net photosynthetic rates on a dry mass basis at both ambient CO(2) (P(n,amb)) and saturating CO(2) (P(n,sat)) than trees in the healthy stand. There were significant linear correlations between P(n,amb) and leaf mass per unit area (LMA) and between P(n,sat) per unit leaf area and LMA. There were also linear correlations between both P(n,amb) and P(n,sat) and leaf N when expressed on an area basis in both stands, indicating that variation in LMA may have been largely responsible for the observed photosynthesis-nitrogen relationship. The values of P(n,amb) and P(n,sat) were not significantly correlated with leaf N on a mass basis but were weakly correlated with leaf Ca and Mg on a mass basis. We conclude that low leaf Ca or Mg concentrations may limit leaf CO(2) assimilation and tree carbohydrate status in the declining stand.     

Development of canopy stratification during early succession in northern hardwoods

Canopy development on a 6-year-old strip cut was analyzed by measuring the heights to terminal buds and bud scale scars of the tallest individuals of each species present on 50 plots of radius 6 m. Phenology of height growth was monitored during the following growing season. Pin cherry (Prunus pensylvania L.), an intolerant short-lived tree, had the fastest growth rate and was on the average the tallest species from the second to the sixth year of regrowth. Although advance regeneration of sugar maple (Acer saccharum Marsh.) and beech (Fagus grandifoloa Ehrh.) were the tallest trees during the first growing season, their slower growth rate insured that they would not keep up with the pin cherry. Trembling aspen (Populus tremuloides Michx.), striped maple (Acer pensylvanicum L.) and yellow birch (Betula alleghaniensis Britt.) occupied an intermediate position in the canopy by the end of the sixth growing season, and showed relatively greater annual height increment than beech or sugar maple. Height growth phenology differed slightly for each species. Beech, ash (Fraxinus americana L.) and sugar maple commenced growth early, grew rapidly and set buds all by 1 August (beech by 15 June). Yellow birch, pin cherry and trembling aspen started growing as early as the others, grew more slowly at first but then grew for a longer period of time. Striped maple seemed to be somewhat intermediate. Growth phenology and growth rate are related to the tolerance and growth form type (e.g. determinate or indeterminate) of the species. The most tolerant species tend to be determinate in growth form, have slower growth rates and complete height growth earlier. The intolerant species tend to be indeterminate, have a faster growth rate and continue to grow for a longer period. These may be mechanisms by which many species can grow together and avoid adverse effects such as suppression.     

Responses of secondary chemicals in sugar maple (Acer saccharum) seedlings to UV-B, springtime warming and nitrogen additions

Anticipated effects of climate change involve complex interactions in the field. To assess the effects of springtime warming, ambient ultraviolet-B radiation (UV-B) and nitrogen fertilization on the foliar chemistry and herbivore activity of native sugar maple (Acer saccharum Marsh.) seedlings, we carried out a field experiment for 2 years at two sugar maple forests growing on soils of contrasting acidity. At the Oliver site, soils are derived from a strongly calcareous till, whereas the naturally acidic soils and base-poor soils of the Haliburton site are derived from the largely granitic Precambrian Shield. At both sites, removal of ambient UV-B led to increases in chlorogenic acid and some flavonoids and reduced herbivore activity. At Haliburton, ammonium nitrate fertilization led to further increases in foliar manganese (Mn), whereas at Oliver there were no such changes. Nitrogen additions led to decreases in the concentrations of some flavonoids at both sites, but seedlings at Oliver had significantly higher concentrations of flavonoids and chlorogenic acid than seedlings at Haliburton. We suggest that this could be associated with increased mobilization of Mn due to increased soil acidity, which interferes with the role of calcium (Ca) in the phenolic biosynthetic pathway. It appears that the composition of the forest soil governs the response of seedlings when they are exposed to abiotic stressors.     

Influence of browsing damage and overstory cover on regeneration of American beech and sugar maple nine years following understory herbicide release in central Maine

Northern hardwood stands, notably those with American beech (Fagus grandifolia Ehrh), sugar maple (Acer saccharum Marsh.), and yellow birch (Betula alleghaniensis Britton), are abundant across the forested landscapes of northeastern USA and southeastern Canada. Recent studies have reported an increasing dominance of American beech in the understory and midstory of these forests. Beech is a commercially less desirable tree species due to its association with beech-bark disease, and because it commonly interferes with the regeneration of other more desirable tree species. We examined hardwood regeneration characteristics nine years after application of a 3 × 4 factorial combination of glyphosate herbicide (0.56, 1.12, and 1.68 kg ha^?1) and surfactant concentrations (0.0, 0.25, 0.5, and 1.0% v v^?1) to release sugar maple regeneration from beech-dominated understories using three stands that received shelterwood seed cutting in central Maine. Measurements nine years after treatment showed that glyphosate rate increased both absolute (AD) and relative density (RD) of sugar maple regeneration, but not its height (HT). In contrast, beech AD, RD, and HT were all significantly reduced with increasing glyphosate rate. Post-release browsing by ungulates and a high residual overstory basal area resulted in reduced sugar maple HT. Our results indicated that glyphosate herbicide applied in stands that have been recently shelterwood seed cut can significantly increase the abundance of sugar maple regeneration. However, subsequent browsing damage combined with the negative influence of the residual overstory cover can limit the longer-term benefit of understory herbicide treatments. Subsequent removal of the overstory and browsing-control measures may be needed to promote sugar maple regeneration over beech in similar northern hardwood stands.     

Leaf phenology,photosynthesis, and the persistence of saplings and shrubs in a mature northern hardwood forest

We quantified leaf phenologies of saplings and overstory trees of sugar maple (Acer saccharum Marsh.) and American beech (Fagus grandifolia Ehrh.), and the shrub hobblebush viburnum (Viburnum alnifolium Marsh.) in a 72-year-old northern hardwood forest. Seasonal changes in irradiance in the shrub layer, and in the leaf CO(2) exchange of viburnum, and sugar maple and beech saplings were also measured. Leaf expansion occurred earlier in the spring and green leaves were retained later in the autumn in saplings and shrubs than in overstory trees. During the spring light phase (before overstory closure), large CO(2) gains by all three shrub-layer species occurred as a result of a combination of relatively large leaf area, high photosynthetic capacity, and high irradiance. Throughout the summer shade phase, photosynthetic capacity at a given irradiance remained relatively constant, but CO(2) gain was typically limited by low irradiances. Even though irradiance in the shrub layer increased during the autumn light phase as the overstory opened, CO(2) gains were modest compared to springtime values because of declining leaf area and photosynthetic capacity in all three species. The CO(2) gains during the spring light phase, and to a lesser extent during the autumn light phase, may be important to the carbon balance and long-term persistence of saplings and shrubs in the usually light-limited shrub layer of a northern hardwood forest. Therefore, for some late-successional species, leaf phenology may be an important characteristic that permits their long-term persistence in the shrub layer of mature northern hardwood forests.     

Evaluation of a portable chlorophyll meter to estimate chlorophyll and nitrogen contents in sugar maple (Acer saccharum Marsh.) leaves

Portable, non-destructive chlorophyll meters could be a valuable tool for forest managers and researchers. The ability of such meters to assess both chlorophyll and nitrogen content has been well established for many agricultural species, but not well-studied for North American forest tree species. The objective of this study was to evaluate the ability of a portable chlorophyll meter to estimate chlorophyll (CHL) and nitrogen (N) content in sugar maple (Acer saccharum Marsh.) leaves by testing correlations between chlorophyll content index values (CCI) and extractable CHL and N in large heterogeneous samples. Significant correlations were observed between CCI and CHL (P ≤ 0.001, r² = 76%) and CCI and N (P ≤ 0.001, r² = 64%). These data indicate that the meter tested can be an effective tool for estimating relative chlorophyll and nitrogen content in sugar maple leaves.     

Neutralization and buffering capacity of leaves of sugar maple, largetooth aspen, paper birch and balsam fir

We compared the acidity, the external acid neutralizing capacity and the buffering capacity of leaves of four commercially important tree species, largetooth aspen (Populus grandidentata Michx.), sugar maple (Acer saccharum Marsh.), paper birch (Betula papyrifera Marsh.) and balsam fir (Abies balsamea (L.) Mill), at two sites of contrasting soil fertility in southern Quebec. External acid neutralizing capacity (ENC) of leaves was determined by measuring the change in pH induced by soaking fresh leaves in an acidic solution (pH 4.0) for two hours. The ENC was highest for largetooth aspen (14.3 micro equiv H(+) g(-1)), and lowest for sugar maple and balsam fir (< 5 micro equiv H(+) g(-1)). The buffering capacity index (BCI) was determined by measuring the amount of acid necessary to produce a change of 5 micro equiv H(+) in the leaf homogenate. The BCI ranged from 883 micro equiv H(+) g(-1) for largetooth aspen to less than 105 micro equiv H(+) g(-1) for sugar maple and balsam fir. Leaves of sugar maple and balsam fir had a lower internal pH and a higher percentage of ENC over BCI than paper birch and largetooth aspen. Overall, ENC was correlated with the concentration of all leaf nutrients except Ca, and BCI was correlated with Mg, N and Ca. The site effect was relatively unimportant for all variables.     

Element accumulation patterns of deciduous and evergreen tree seedlings on acid soils: implications for sensitivity to manganese toxicity

Foliar nutrient imbalances, including the hyperaccumulation of manganese (Mn), are correlated with symptoms of declining health in sensitive tree species growing on acidic forest soils. The objectives of this study were to: (1) compare foliar nutrient accumulation patterns of six deciduous (sugar maple (Acer saccharum Marsh.), red maple (Acer rubrum L.), red oak (Quercus rubra L.), white oak (Quercus alba L.), black cherry (Prunus serotina Ehrh.) and white ash (Fraxinus americana L.)) and three evergreen (eastern hemlock (Tsuga canadensis L.), white pine (Pinus strobus L.) and white spruce (Picea glauca (Moench) Voss.)) tree species growing on acidic forest soils; and (2) examine how leaf phenology and other traits that distinguish evergreen and deciduous tree species influence foliar Mn accumulation rates and sensitivity to excess Mn. For the first objective, leaf samples of seedlings from five acidic, non-glaciated field sites on Pennsylvania's Allegheny Plateau were collected and analyzed for leaf element concentrations. In a second study, we examined growth and photosynthetic responses of seedlings exposed to excess Mn in sand culture. In field samples, Mn in deciduous foliage hyperaccumulated to concentrations more than twice as high as those found in evergreen needles. Among species, sugar maple was the most sensitive to excess Mn based on growth and photosynthetic measurements. Photosynthesis in red maple and red oak was also sensitive to excess Mn, whereas white oak, black cherry, white ash and the three evergreen species were tolerant of excess Mn. Among the nine species, relative rates of photosynthesis were negatively correlated with foliar Mn concentrations, suggesting that photosynthetic sensitivity to Mn is a function of its rate of accumulation in seedling foliage.     

Growth,chlorophyll fluorescence and biochemical markers in clonal ramets of shisham (<Emphasis Type="Italic">Dalbergia sissoo</Emphasis> Roxb.) at nursery stage

About 19 clones of shisham (Dalbergia sissoo Roxb.) were raised by using healthy coppice shoot cuttings. After 2 year of establishment and growth, they were examined to identify suitable clone of shisham at nursery stage. Clones were studied for growth (height and basal diameter), chlorophyll fluorescence (Fv/Fm) and biochemical contents (chlorophyll a, b, total chlorophyll, free amino acids, total soluble proteins, total soluble sugars, starch and reducing sugar) in the leaves. Clone C2 and C3 (Gonda, Uttar Pradesh, India) had the highest growth and the chlorophyll fluorescence rate among examined clones. Between these two clones, C2 gave the highest response on height, basal diameter and Fv/Fm. Except free amino acids, all other biochemical contents were greater in the clones belonging to Gonda, Uttar Pradesh, India. Growth parameters, i.e. height and basal diameter increment at nursery stage had a positive correlation with all the studied parameters, except free amino acid and total soluble protein. These findings have demonstrated that there are remarkable clonal variations in growth, physiological and biochemical responses at nursery stage. Thus, physiological and biochemical markers can enhance selection efficiency, in addition to growth characteristics such as height and basal diameter as these have practical value in the tree improvement. The study suggests that clones of Gonda, Uttar Pradesh, India had the best performance and could be further tested in plantation programs.     

Seasonal changes of C and N non-structural compounds in the stem sapwood of adult sessile oak and beech trees

We assessed the pools of non-structural nitrogen compounds (NSNC) through a year, thereby addressing the question of whether mature sessile oak [Quercus petraea (Matt.) Liebl.] and beech (Fagus sylvatica L.), which differ in wood anatomy and growth patterns, exhibit contrasting seasonal dynamics of NSNC pools as previously shown for non-structural carbohydrate (NSC) pools. Seasonal fluctuations of NSNC (amino acids and soluble proteins) and NSC (starch and soluble sugars) pools were analyzed in the inner and the outer stem sapwood. In oak, NSC showed marked seasonal variation within the stem sapwood (accumulation during winter and decrease during bud burst and early wood growth), whereas in beech seasonal fluctuations in NSC were of minor amplitude. Even if the distribution and intensity of the NSNC pools differed between the two species, NSNC of the stem sapwood did not show seasonal variation. The most significant change in NSNC pools was the seasonal fluctuation of protein composition. In both species, two polypeptides of 13 kDa (PP13) and 26 kDa (PP26) accumulated during the coldest period in parallel with starch to sugar conversion and disappeared with the onset of spring growth. The absence of seasonal changes in total soluble protein concentration suggests that the polypeptides are involved in the internal nitrogen (N) cycling of the stem rather than in N storage and remobilization to the other growing organs of the tree.     

Influence of nitrogen fertilization on minerals, carbohydrates, amino acids and phenolic compounds in beech (Fagus sylvatica L.) leaves

Beginning in 1985, plots in a 120-year-old beech (Fagus sylvatica L.) stand in the province of Scania, southern Sweden were fertilized annually with ammonium nitrate. In August 1989, leaves from both fertilized and unfertilized trees were analyzed for mineral nutrients, soluble carbohydrates, amino acids and phenolic compounds. Leaf concentrations of total nitrogen were increased by N fertilization. Ammonium was not detected in the leaves, but NO(3) (-) was increased by up to 65% as a result of N fertilization. Nitrogen fertilization decreased foliar concentrations of P and Cu, leading to a nutrient imbalance relative to N. Of the free amino acids, glutamic acid and aspartic acid predominated and, together with glutamine and asparagine, were significantly increased by N fertilization. The concentrations of amides were 2-4 times higher in fertilized trees than in unfertilized trees. Soluble carbohydrates did not change in response to N fertilization, but total phenolic compounds decreased markedly. The reduced concentration of phenolics may explain the greater susceptibility to parasite attacks of trees exposed to N deposition and excess N.     

Factors influencing red expression in autumn foliage of sugar maple trees

We evaluated factors influencing the development of autumn red coloration in leaves of sugar maple (Acer saccharum Marsh.) by measuring mineral nutrient and carbohydrate concentrations, water content, and phenology of color development of leaves from 16 mature open-grown trees on 12 dates from June through October 1999. Mean foliar nutrient and carbohydrate concentrations and water content were generally within the range published for healthy sugar maple trees. However, foliar nitrogen (N) concentrations were near deficiency values for some trees. The timing and extent of red leaf coloration was consistently correlated with both foliar N concentrations and starch or sugar concentrations, which also varied with N status. Leaves of trees with low foliar N concentrations turned red earlier and more completely than those of trees with high foliar N concentrations. Low-N trees also had higher foliar starch concentrations than high-N trees. During the autumn development of red leaf coloration, foliar starch, glucose and fructose concentrations were positively correlated with red leaf color expression. At peak red expression, the concentrations of glucose, fructose, sucrose and stachyose were all positively correlated with red color expressed as a percent of total leaf area.