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.     

Estimating water use by sugar maple trees: considerations when using heat-pulse methods in trees with deep functional sapwood

Accurate estimates of sapwood properties (including radial depth of functional xylem and wood water content) are critical when using the heat pulse velocity (HPV) technique to estimate tree water use. Errors in estimating the volumetric water content (V(h)) of the sapwood, especially in tree species with a large proportion of sapwood, can cause significant errors in the calculations ofsap velocity and sap flow through tree boles. Scaling to the whole-stand level greatly inflates these errors. We determined the effects of season, tree size and radial wood depth on V(h) of wood cores removed from Acer saccharum Marsh. trees throughout 3 years in upstate New York. We also determined the effects of variation in V(h) on sap velocity and sap flow calculations based on HPV data collected from sap flow gauges inserted at four depths. In addition, we compared two modifications of Hatton's weighted average technique, the zero-step and zero-average methods, for determining sap velocity and sap flow at depths beyond those penetrated by the sap flow gauges. Parameter V(h) varied significantly with time of year (DOY), tree size (S), and radial wood depth (RD), and there were significant DOY x S and DOY x RD interactions. Use of a mean whole-tree V(h) value resulted in differences ranging from -6 to +47% for both sap velocity and sap flow for individual sapwood annuli compared with use of the V(h) value determined at the specific depth where a probe was placed. Whole-tree sap flow was 7% higher when calculated on the basis of the individual V(h) value compared with the mean whole-tree V(h) value. Calculated total sap flow for a tree with a DBH of 48.8 cm was 13 and 19% less using the zero-step and the zero-average velocity techniques, respectively, than the value obtained with Hatton's weighted average technique. Smaller differences among the three methods were observed for a tree with a DBH of 24.4 cm. We conclude that, for Acer saccharum: (1) mean V(h) changes significantly during the year and can range from nearly 50% during winter and early spring, to 20% during the growing season;(2) large trees have a significantly greater V(h) than small trees; (3) overall, V(h) decreases and then increases significantly with radial wood depth, suggesting that radial water movement and storage are highly dynamic; and (4) V(h) estimates can vary greatly and influence subsequent water use calculations depending on whether an average or an individual V(h) value for a wood core is used. For large diameter trees in which sapwood comprises a large fraction of total stem cross-sectional area (where sap flow gauges cannot be inserted across the entire cross-sectional area), the zero-average modification of Hatton's weighted average method reduces the potential for large errors in whole-tree and landscape water balance estimates based on the HPV method.     

Individual-tree diameter growth model for sugar maple trees in uneven-aged northern hardwood stands under selection system

An individual-tree diameter model was developed for sugar maple (Acer saccharum Marsh.) in northern hardwood stands managed under selection system. We fitted long-term remeasurement data to a linear mixed model to account for the temporal autocorrelation of the remeasurements. The model was evaluated using independent data from two physiographic regions and representing a range of tree diameter classes, residual basal areas and years since cut. We compared our model to several individual-tree models based on data from stands with varied management histories. Several competition indices were also tested for an improvement in model fitting and prediction. Our model had lower bias and prediction error when compared to two previous models, as it better accounted for the increased diameter growth that occurred in trees from appropriately managed stands. The addition of a tree-specific competition index failed to improve model fit and predictive ability over stand-level basal area.     

Frost tolerance and bud dormancy of container-grown yellow birch, red oak and sugar maple seedlings

Container-grown seedlings of red oak (Quercus rubra L.), sugar maple (Acer saccharum Marsh.) and yellow birch (Betula alleghaniensis Britton) in their first year of growth were overwintered outdoors. Tolerance of roots and stems to freezing was compared from late summer to the following spring. Mitotic activity in the apical bud was related more closely to air temperature than to bud dormancy as defined by days to bud break. In all species, stem hardening was observed before days to bud break reached a maximum. Dormancy release (days to bud break equal to zero) of yellow birch coincided with loss of stem hardening in the spring. Roots hardened more slowly, had a lower frost tolerance than stems in fall and winter, and dehardened earlier than stems in the spring. There were differences in stem and root hardiness among the species, with yellow birch being the most tolerant, followed by sugar maple and red oak. Primarily because of root sensitivity to frost, winter was a critical period for all three species, but particularly for red oak.     

Productivity and growth efficiency in sugar maple forests

Sugar maple (Acer saccharum Marsh.)-dominated northern hardwood forests were examined at four locations along an acid deposition and climatic gradient in the Great Lakes region of the USA. The study sites were matched in terms of physiography, soils, stand history, and vegetative characteristics. Measurements of basal area and biomass growth were made for the 1988–1991 growing seasons. There were no significant differences in either basal area of biomass increment among the four sites over the 4 year period. There was a great deal of year-to-year variability with relative basal area growth rates ranging from as low as 0.2% to as high as 2.4% on a single site in successive years. Growth efficiency measures reflected this variability with as much as an 800% difference between successive years on a single site. When coupled with year-to-year variability of up to 34% in leaf area related to heavy seed years and defoliation, this indicates that growth efficiency and leaf area measures are not consistent indicators of aboveground productivity for tolerant deciduous species, especially if derived from short-term measurements or temporary plots.     

Field measurements of root respiration indicate little to no seasonal temperature acclimation for sugar maple and red pine

Increasing global temperatures could potentially cause large increases in root respiration and associated soil CO2 efflux. However, if root respiration acclimates to higher temperatures, increases in soil CO2 efflux from this source would be much less. Throughout the snow-free season, we measured fine root respiration in the field at ambient soil temperature in a sugar maple (Acer saccharum Marsh.) forest and a red pine (Pinus resinosa Ait.) plantation in Michigan. The objectives were to determine effects of soil temperature, soil water availability and experimental N additions on root respiration rates, and to test for temperature acclimation in response to seasonal changes in soil temperature. Soil temperature and soil water availability were important predictors of root respiration and together explained 76% of the variation in root respiration rates in the red pine plantation and 71% of the variation in the sugar maple forest. Root N concentration explained an additional 6% of the variation in the sugar maple trees. Experimental N additions did not affect root respiration rates at either site. From April to November, root respiration rates measured in the field increased exponentially with increasing soil temperature. For sugar maple, long-term Q10 values calculated from the field data were slightly, but not significantly, less than short-term Q10 values determined for instantaneous temperature series conducted in the laboratory (2.4 versus 2.62.7). For red pine, long-term and short-term Q10 values were similar (3.0 versus 3.0). Sugar maple root respiration rates at constant reference temperatures of 6, 18 and 24 degrees C were measured in the laboratory at various times during the year when field soil temperatures varied from 0.4 to 16.8 degrees C. No relationship existed between ambient soil temperature just before sampling and root respiration rates at 6 and 18 degrees C (P = 0.37 and 0.86, respectively), and only a very weak relationship was found between ambient soil temperature and root respiration at 24 degrees C (P = 0.08, slope = 0.09). We conclude that root respiration in these species undergoes little, if any, acclimation to seasonal changes in soil temperature.     

Influence of ozone on cold acclimation in sugar maple seedlings

During summer 1994, sugar maple (Acer saccharum Marsh.) seedlings were grown in open-top chambers supplied with air containing near ambient ozone concentration (control, low O(3)) or three times the ambient ozone concentration (high O(3)). The rate of CO(2) assimilation was significantly reduced by chronic exposure to a high concentration of ozone during the summer. During fall, seedlings were removed from the open-top chambers and acclimated to cold under natural conditions. In both species during cold acclimation, the starch concentration decreased, whereas the sucrose concentration increased. There was no treatment effect on the freezing tolerance of roots, even though roots in the high-O(3) treatment accumulated higher concentrations of the cryoprotective oligosaccharides raffinose and stachyose than control roots. Cold acclimation occurred earlier and stachyose concentration of stems was higher in high-O(3)-treated seedlings than in low-O(3)-treated seedlings. Cold acclimation was associated with an earlier accumulation of ABA in the xylem sap of high-O(3)-treated seedlings compared with low-O(3)-treated seedlings.     

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.     

Carbon content variation in boles of mature sugar maple and giant sequoia

At present, a carbon (C) content of 50% (w/w) in dry wood is widely accepted as a generic value; however, few wood C measurements have been reported. We used elemental analysis to investigate C content per unit of dry matter and observed that it varied both radially and vertically in boles of two old-growth tree species: sugar maple (Acer saccharum Marsh.) and giant sequoia (Sequoiadendron giganteum (Lindl.) Bucholz). In sugar maple there was considerable variation in tree ring widths among four radii for particular annual layers of xylem, revealing that the annual rate of C assimilation differs around the circumference and from the base of each tree to its top, but the observed variation in C content was unrelated to diameter growth rate and strongly related to the calendar year when the wood was formed. Carbon content in sugar maple wood increased in an approximately linear fashion, from < 50 to 51% from pith to cambium, at both the base and top of the boles. In giant sequoia, C was essentially constant at > 55% across many hundreds of years of heartwood, but it declined abruptly at the sapwood-heartwood boundary and remained lower in all sapwood samples, an indication that heartwood formation involves anabolic metabolism. Factors that may be responsible for the different C contents and trends with age between sugar maple and sequoia trees are considered. Tree-ring data from this study do not support some of the key assumptions made by dendrochronology.     

The successional status of sugar maple (Acer saccharum), revisited

Two complementary experimental designs at two contrasting scales (landscape/long term; individual tree/short term) were used for an in-depth evaluation of the successional status of sugar maple (AS: Acer saccharum Marsh.). First, forest disturbances during the 20th century and composition were mapped for two landscapes in the Du Lièvre watershed of southern Quebec. Our results show that, as well as dominating stands in the absence of fire, AS often rapidly developed dominance after fire, especially in the south of our study area. Similarly, a majority of AS-dominated stands clearcut in 1928 continued to be AS-dominated 60 years later. Second, we examined AS seedlings planted under two very contrasting light regimes. AS seedlings showed a combination of traits particularly adapted to tolerate shade under a low light regime. However, owing to a surprisingly high phenotypic plasticity, AS also exhibited efficient development under high light. This suggests the classification of AS as a late-successional species should indeed be revised and that generalist or trans-successional would be a more appropriate designation for this species. We discuss the ramifications of such a status revision, with an emphasis on the implications for its silviculture.     

Variation in sugar maple root respiration with root diameter and soil depth

Root respiration may account for as much as 60% of total soil respiration. Therefore, factors that regulate the metabolic activity of roots and associated microbes are an important component of terrestrial carbon budgets. Root systems are often sampled by diameter and depth classes to enable researchers to process samples in a systematic and timely fashion. We recently discovered that small, lateral roots at the distal end of the root system have much greater tissue N concentrations than larger roots, and this led to the hypothesis that the smallest roots have significantly higher rates of respiration than larger roots. This study was designed to determine if root respiration is related to root diameter or the location of roots in the soil profile. We examined relationships among root respiration rates and N concentration in four diameter classes from three soil depths in two sugar maple (Acer saccharum Marsh.) forests in Michigan. Root respiration declined as root diameter increased and was lower at deeper soil depths than at the soil surface. Surface roots (0-10 cm depth) respired at rates up to 40% greater than deeper roots, and respiration rates for roots < 0.5 mm in diameter were 2.4 to 3.4 times higher than those for roots in larger diameter classes. Root N concentration explained 70% of the observed variation in respiration across sites and size and depth classes. Differences in respiration among root diameter classes and soil depths appeared to be consistent with hypothesized effects of variation in root function on metabolic activity. Among roots, very fine roots in zones of high nutrient availability had the highest respiration rates. Large roots and roots from depths of low nutrient availability had low respiration rates consistent with structural and transport functions rather than with active nutrient uptake and assimilation. These results suggest that broadly defined root classes, e.g., fine roots are equivalent to all roots < 2.0 mm in diameter, do not accurately reflect the functional categories typically associated with fine roots. Tissue N concentration or N content (mass x concentration N) may be a better indicator of root function than root diameter.     

Rapid freezing induces winter injury symptomatology in red spruce foliage

Red spruce (Picea rubens Sarg.) suffers frequent and extensive injury to current-year foliage during the winter. Experimental freezing of red spruce foliage at cooling rates > 10 degrees C min(-1) induced visible symptomatology similar to natural winter injury at the branch, needle and cellular levels. Such damage was associated with a low-temperature exotherm near -10 to -12 degrees C, a loss in needle fluorescence, massive cellular disruption, foliar discoloration, and low needle survival. Susceptibility of individual trees to rapid freezing injury was associated with historical winter injury patterns and alterations in foliar nutrition. We conclude that anthropogenic deposition may alter the sensitivity of trees to winter injury caused by rapid temperature changes.     

Comparison analysis of nonlinear segmentation methods for selected wood features in sugar maple

Two classification methods, a feed-forward neural network and a fuzzy logic algorithm, were used for the automatic identification of CT images for selected wood features in sugar maple, one of the most important hardwoods in eastern Canada. Three wood characteristics were selected for automatic identification together with the background as a default. Local features, such as position and local pixel values were used as the neural networks inputs. The fuzzy sets consisted of four different possible pixel values and four possible distances from the center of the log. The fuzzy method used in this study was of the Mamdani type. Five sugar maple logs were randomly selected for this study. One of the logs is used for the training of the neural network and the others for validation and comparison. The structure of the neural network was optimized and was used for the segmentation of the other logs. An efficiency function, consisting of the number of pixels correctly labeled, was defined for the evaluation of the segmentation process. This study shows that a segmentation based on a fuzzy method has better capabilities for generalization than one based on a feed-forward method.     

Stress wave sorting of red maple logs for structural quality

Existing log grading procedures in the United States make only visual assessments of log quality. These procedures do not incorporate estimates of the modulus of elasticity (MOE) of logs. It is questionable whether the visual grading procedures currently used for logs adequately assess the potential quality of structural products manufactured from them, especially those for which MOE is of primary concern. The purpose of this study was to investigate the use of stress wave nondestructive evaluation techniques to sort red maple logs for the potential quality of lumber obtained from them. Ninety-five red maple logs were nondestructively evaluated using longitudinal stress wave techniques and sorted into four stress wave grades. The logs were then sawn into cants and lumber. The same procedure was used to obtain stress wave times in the cants and lumber. The lumber specimens were then dried and graded using a transverse vibration technique. The results of this study showed that good relationships existed between stress wave times measured in logs, cants, and the lumber produced from the logs. It was found that log stress wave grades have positive relationships with the lumber grades. Logs with high stress wave grades produced high-grade lumber. These findings indicate that the longitudinal stress wave technique has potential in sorting logs and cants for the production of high MOE products.The Forest Products Laboratory is maintained in cooperation with the University of Wisconsin. This article was written and prepared by U.S. Government employees on official time, and it is therefore in the public domain and not subject to copyright.     

Electrical resistance and stem water relations in Verticillium-infected red maple1

An inverse relationship exists between stem water potential and electrical resistance (ER) in healthy red maple Acer rubrum L. (r = - .93 to r = - .98). Stem ER of red maple seedlings was measured at 1 to 2 day intervals during a 14 week course of infection by Verticillium dahliae Klebahn. No significant differences (P = .05) in ER were found between moderately to severely infected and control plants, although stem water potential of infected plants was significantly lower than the control plants. In plants which displayed only slight if any symptoms and in which no significant change in stem water potential occurred, ER of the infected plants was significantly lower than that of the control plants.     

Influence of wood planing on the second-order effects of moisture sorption in sugar maple

Summary Two types of machines, a conventional planer, and a fixed-knife pressure-bar planer were used to prepare matched specimens of sugar maple wood. After adsorption and desorption, both experiments at 21 °C, the EMC, swelling in all principal directions as well as compliance coefficient in radial compression were measured. Two specimen sizes were used for these expe‐riments. For a given equilibrium moisture content, tangential and radial dimensions were greater after desorption than after adsorption, as previously described. When equilibrium was reached by gaining moisture, the wood was stiffer in radial compression compared to when the equilibrium was reached after losing moisture. The magnitude of this phenomenon, second-order effects of moisture sorption, was slightly affected by the type of planing. These effects on swelling were greater for large specimens prepared by conventional planing compared to fixed-knife pressure-bar planing. Small specimens showed similar magnitudes of this phenomenon with both planing methods. No differences between planing methods were found for the radial compliance coefficient measured on either specimen size. Therefore, the second-order effects of moisture sorption appeared to be a bulk phenomenon and not restricted to the superficial layers of wood. Received 9 December 1997     

Influence of overstory density on ecophysiology of red oak (Quercus rubra) and sugar maple (Acer saccharum) seedlings in central Ontario shelterwoods

A field experiment was established in a second-growth hardwood forest dominated by red oak (Quercus rubra L.) to examine the effects of shelterwood overstory density on leaf gas exchange and seedling water status of planted red oak, naturally regenerated red oak and sugar maple (Acer saccharum Marsh.) seedlings during the first growing season following harvest. Canopy cover of uncut control stands and moderate and light shelterwoods averaged 97, 80 and 49%, respectively. Understory light and vapor pressure deficit (VPD) strongly influenced gas exchange responses to overstory reduction. Increased irradiance beneath the shelterwoods significantly increased net photosynthesis (P(n)) and leaf conductance to water vapor (G(wv)) of red oak and maple seedlings; however, P(n) and G(wv) of planted and naturally regenerated red oak seedlings were two to three times higher than those of sugar maple seedlings in both partial harvest treatments, due in large part to decreased stomatal limitation of gas exchange in red oak as a result of increased VPD in the shelterwoods. In both species, seedling water status was higher in the partial harvest treatments, as reflected by the higher predawn leaf water potential and seedling water-use efficiency in seedlings in shelterwoods than in uncut stands. Within a treatment, planted and natural red oak seedlings exhibited similar leaf gas exchange rates and water status, indicating little adverse physiological effect of transplanting. We conclude that the use of shelterwoods favors photosynthetic potential of red oak over sugar maple, and should improve red oak regeneration in Ontario.     

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.