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.     

Root carbohydrate reserves, mineral nutrient concentrations and biomass in a healthy and a declining sugar maple (Acer saccharum) stand

Soil and root characteristics were contrasted between a "declining" and a "healthy" sugar maple (Acer saccharum Marsh.) stand in Vermont, USA. The declining stand had lower basal area increment and more crown dieback than the healthy stand. Soil pH and base cation content were lower and soil water content was higher at the site of the declining stand than at the site of the healthy stand, whereas soil temperature did not differ significantly between the sites. In live fine roots, concentrations of K and Ca were marginally (P < 0.07) lower in the declining than in the healthy stand, whereas concentrations of N, P, Mg, and Al were not significantly different (P = 0.13 to 0.87) between stands. Starch and soluble sugar concentrations of fine and coarse roots did not differ significantly between stands, indicating that crown dieback did not affect carbohydrate supply to the roots in the declining stand. Throughout the growing season, the standing live and dead root biomass were significantly higher in the declining stand than in the healthy stand, indicating that more carbon was allocated to roots and that root turnover was higher in the declining stand than in the healthy stand.     

Micro-variations in yellow birch (Betula alleghaniensis) growth conditions after patch scarification

The abundance of poor quality yellow birch stands resulting from diameter limit cuts poses important regeneration challenges since seed production is uncertain, adequate seedbeds are rare and competition very severe. The present study aims at evaluating variations in seedling growth, temperature, soil humidity and nutrient availability for the whole range of seedbeds created by a patch scarification treatment. In three poor quality stands, soil temperature and humidity, nutrient availability and seedling morphology were monitored on undisturbed forest floor, at different places in 2 m wide scarified patches, and on mounds formed by the displaced material. Even though the study associated some variations in temperature and humidity to different microsite positions, the effect on nutrient availability was more pronounced. Hence, the study found better nutrient availability on mounds in comparison with patches as well as at the border of patches. Soil nutrient availability as measured with in situ ion exchange resin declined very rapidly in the interior of scarified patch (within 5 cm for the border) suggesting that small scarified patches combined with control of vegetation competition should provide a better environment for seedling growth. Growth parameters of yellow birch seedlings were not correlated to measured soil parameters suggesting that there were important effects of uncontrolled factors.     

In vivo and in situ rhizosphere respiration in Acer saccharum and Betula alleghaniensis seedlings grown in contrasting light regimes

A perfusive method combined with an open-system carbon dioxide measurement system was used to assess rhizosphere respiration of Acer saccharum Marsh. (sugar maple) and Betula alleghaniensis Britton (yellow birch) seedlings grown in 8-l pots filled with coarse sand. We compared in vivo and in situ rhizosphere respiration between species, among light regimes (40, 17 and 6% of full daylight) and at different times during the day. To compute specific rhizosphere respiration, temperature corrections were made with either species-specific coefficients (Q10) based on the observed change in respiration rate between 15 and 21 degrees C or an arbitrarily assigned Q10 of 2. Estimated, species-specific Q10 values were 3.0 and 3.4 for A. saccharum and B. alleghaniensis, respectively, and did not vary with light regime. Using either method of temperature correction, specific rhizosphere respiration did not differ either between A. saccharum and B. alleghaniensis, or among light regimes except in A. saccharum at 6% of full daylight. At this irradiance, seedlings were smaller than in the other light treatments, with a larger fine root fraction of total root dry mass, resulting in higher respiration rates. Specific rhizosphere respiration was significantly higher during the afternoon than at other times of day when temperature-corrected on the basis of an arbitrary Q10 of 2, suggesting the possibility of diurnal variation in a temperature-independent component of rhizosphere respiration.     

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.     

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.     

Structural differences and functional similarities between two sugar maple (Acer saccharum) stands

The spatially inexplicit or functional multilayer models used to predict canopy transpiration or photosynthesis are based on the assumption that closed stands show less functional variability than structural variability, because foliage tends to arrange itself in space to optimize the capture of light. To validate this assumption, we compared the structural and functional properties, and the measured and modeled transpiration fluxes of two sugar maple (Acer saccharum Marsh.) stands of comparable leaf mass but differing in height and diameter distributions. One stand was characterized by a well-developed single-layer canopy, whereas the other stand had a multilayered canopy and a stem diameter distribution of the classical inverse-J shape. Stand differences in height and diameter distribution, and canopy gap fraction, were highly significant. There were minor but significant differences in leaf mass and leaf mass per unit leaf area (LMA) distributions. We found no differences in tree-level relationships between basal area and either transpiration flux or sapwood area. We compared measurements of stand transpiration with transpiration estimates obtained from a multilayer gas exchange model, in which only the nonspatial inputs, leaf area index and LMA frequency distribution described stand structure. For both stands, modeled values of daily transpiration closely followed measured values (r(2) = 0.94). These results support use of the nonspatially explicit approach to estimating canopy gas exchange, especially if the intent is to scale-up to larger portions of the landscape.     

Photosynthesis and water-use efficiency of sugar maple (Acer saccharum) in relation to pear thrips defoliation

An experimental introduction of pear thrips (Taeniothrips inconsequens Uzel), a major defoliator in sugar maple (Acer saccharum Marsh.) forests in northeastern North America, was conducted in a field plantation to determine if compensatory gas exchange occurs in response to feeding damage by this piercing-sucking insect. Sugar maple trees were enclosed in netting (167 micro m mesh) and pear thrips adults were introduced before leaf expansion in the spring. Pear thrips reduced whole-tree leaf area by approximately 23% and reduced leaf size (both mass and area) by 20% in the upper crown. Measurements of net CO(2) assimilation rate (A(net)) and stomatal conductance (g(s)) were made on tagged foliage that was later analyzed for stable carbon isotope composition (delta(13)C) to provide estimates of short- and long-term leaf water use efficiency (WUE). Pear thrips feeding reduced A(net) for fully expanded leaves by approximately 20%, although leaf chlorophyll content and leaf mass per unit area were apparently not affected. Comparison of A(net), g(s), instantaneous WUE and leaf delta(13)C between damaged and control trees as well as visibly undamaged versus moderately damaged foliage on pear thrips-infested trees indicated that there were no effects of pear thrips feeding damage on WUE or leaf delta(13)C. Long-term WUE among sugar maple trees in the field plantation, indicated by leaf delta(13)C analysis, was related to shorter-term estimates of leaf gas exchange behavior such as g(s) and calculated leaf intercellular CO(2) concentration (C(i)). We conclude that pear thrips feeding has no effect on leaf WUE, but at the defoliation levels in our experiment, it may reduce leaf A(net), as a result of direct tissue damage or through reduced g(s). Therefore, even small reductions in leaf A(net) by pear thrips feeding damage may have an important effect on the seasonal carbon balance of sugar maple when integrated over the entire growing season.     

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.     

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.     

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.     

Crown architecture of stand-grown sugar maple (Acer saccharum Marsh.) in the Adirondack Mountains

Leaf and crown morphology of shade-tolerant sugar maple (Acer saccharum Marsh.) were examined to test the hypotheses (1) that leaf area exhibits significant plasticity both within and between crown classes and individual tree crowns and (2) that leaf area is accurately predicted from estimates of crown volume. A total of 18 trees, ranging from 3.3 to 43.4 cm dbh, were felled and dissected into upper, middle, lower, and below-crown layers, for measurements of leaf, bark, and xylem dimensions. For dominant trees only, bark thickness and xylem radii were higher within the crown than below the crown. Cumulative leaf area index increased with decreasing stratum height at similar rates in all trees, except for two trees that were located in the understory. Area leaf weight declined with decreasing stratum height within the crown of all except four overstory trees. These four trees showed an increase with decreasing stratum height, i.e., leaves were heavier per unit area in the lower crown stratum and below the crown than they were at mid-crown. Within-tree leaf area density was usually higher in the upper crown of overstory trees and in the lower crown of understory trees. Total crown volume was the best predictor of whole-tree leaf area, but it was only slightly better than dbh.     

Canopy photosynthesis of sugar maple (Acer saccharum): comparing big-leaf and multilayer extrapolations of leaf-level measurements

A comparison is made between a big-leaf model (i.e., without details of the canopy profile) and two multilayer models (i.e., with details of the canopy profile) to estimate daily canopy photosynthesis of a sugar maple (Acer saccharum Marsh.) stand. The first multilayer model uses the distribution of leaf area by leaf mass per unit area (LMA) classes, the observed relationships between the parameters of a photosynthesis-irradiance curve and LMA, and the relationship between relative irradiance and LMA to estimate canopy photosynthesis. When compared with this model, the big-leaf model underestimates daily canopy photosynthesis by 26% because of an assumed proportionality between photosynthetic capacity and relative irradiance, a proportionality that is inconsistent with our data. The bias induced by this assumption is reduced when the big-leaf model is compared with the second multilayer model which, in addition to the assumptions made for the first multilayer model, accounts for the sunlit and shaded fractions of leaf area. The residual bias is almost eliminated when the big-leaf model is run using a weekly averaged irradiance. It is likely, however, that this is the result of a compensating bias that, in this particular case, compensates for the initial bias introduced by the proportionality assumption. It is also shown that canopy photosynthesis can be represented by spatially inexplicit multilayer models that use leaf mass per area as a covariable to describe leaf characteristics and environment. Such models represent an interesting alternative to the biased big-leaf approach.     

Sticker stain formation in hardwoods: Isolation of scopoletin from sugar maple (Acer saccharum Marsh.)

Summary The extractives of clear and sticker stained sapwood from sugar maple (Acer saccharum Marsh.) were isolated and screened for low molecular-weight phenols, which could be involved in the formation of sticker stain. Scopoletin (7-hydroxy-6-methoxycoumarin) was identified as the major low molecular-weight free phenol in the samples studied. This compound, which has not previously been reported in extractives from maple wood, was quantified in stained and clear samples. Additionally, the two major fatty acids present were identified as palmitic acid and linolenic acid, and the two major sterols as stigmasterol and sitosterol.The authors are indebted to Mr. Peter Garrahan for provision of wood samples and the Canadian Forestry Service for financial support     

Water vapour diffusion and adsorption characteristics of sugar maple (Acer saccharum,Marsh.) wood polymer composites

Summary Water vapour diffusion characteristics and adsorption isotherms were determined for cell-lumen and cell-wall treated wood polymer composites (WPC). The diffusion coefficients of the cell-lumen WPC were lower than untreated wood and the cell-wall WPC coefficients were lower than cell-lumen. Using the Hailwood and Horrobin sorption model, it was found that the unimolecular layer is formed at lower moisture contents in WPC than in wood. The amount of free dissolved water was reduced only in the cell-wall WPC. The polymer reduces the water vapour accessibility in both types of WPC.     

Effects of xylem cavitation and freezing injury on dieback of yellow birch (Betula alleghaniensis) in relation to a simulated winter thaw

Shoot dieback, shoot growth, stem xylem cavitation, stem and root freezing injury, and root pressure were measured in 2-year-old, cold-hardened, potted yellow birch (Betula alleghaniensis Britt.) seedlings that had been subjected to a simulated winter thaw for 0, 5, 10, 19 or 27 days followed by 10 weeks at -10 degrees C. Stem xylem cavitation was determined as percent loss of hydraulic conductivity. Stem freezing injury was measured as electrolyte leakage (EL). Root freezing injury was determined by EL and by triphenyl tetrazolium chloride (TTC) reduction. Thaw duration was significantly correlated with dieback, new shoot growth, stem xylem cavitation, stem and root freezing damage, and root pressure (P < 0.05). In particular, shoot dieback was positively correlated with stem xylem cavitation (P < 0.001), residual stem xylem cavitation (P < 0.01) and root freezing injury (P < 0.010), but only weakly correlated with stem freezing damage (P < 0.05). In roots, freezing damage was negatively correlated with root pressure (P < 0.05), which, in turn, was negatively correlated with residual stem xylem cavitation after root pressure development. In stems, there was no correlation between freezing damage and xylem cavitation. We conclude that long periods of winter thaw followed by freezing resulted in freezing injury to roots concomitant with a reduction in root pressures, leading to poor recovery from freezing-induced xylem embolism.     

Effects of soil compaction on growth variables in Cappadocian maple(Acer cappadocicum) seedlings

This study investigates the effects of increasing soil penetration resistance(SPR) on seedling morphology and seedling architecture. When seedlings of deciduous Cappadocian maple(Acer cappadocicum Gled.) were grown in a greenhouse in a loamy soil under a wide range of soil compactions, all morphological variables studied changed significantly with increasing SPR. The relationships between increasing SPR and all morphological responses except lateral root length followed a negative quadratic curve. All biomass variables except lateral root biomass showed a bell-shaped response with respect to SPR, with a maximum biomass variable between 0.6 and1.2 MPa, decreasing at higher soil compaction values. All allocation ratios were significantly affected by soil penetration resistance. Biomass allocation to roots was also affected by soil compaction. There was not a significant relationship between the specific stem length and increasing soil penetration resistance. The specific root length showed two trends to increasing SPR; it first decreased in response to the moderate compaction treatment(up to about 1.2 MPa), then increased significantly. We concluded that increasing soil compaction caused morphological changes to root and shoot sections of A.cappadocicum seedlings.     

Seasonal and within-stem variations of neutral lipids in silver birch (Betula pendula) wood

Neutral lipids were analyzed in stem wood of a 7-year-old clone and in five 35-70-year-old mature trees of silver birch (Betula pendula Roth). In young trees and in mature wood of old trees, the free fatty acid fraction comprised less than 5% of the concentration of triacylglycerols (TG). The concentration of free linoleic acid was lowest in March when the young trees were dormant and highest during midsummer and September. In mature trees, the TG concentration increased toward the pith, indicating that living parenchyma cells close to the pith have a large TG storage capacity. The TG concentration (mean 0.51 +/- 0.02% of wood dry mass) remained constant throughout the year in young trees, whereas the concentration of beta-sitosterol, the dominant free sterol (mean 82.5 +/- 0.4% of total free sterols), decreased during spring and early summer when the temperature gradually increased, and increased during autumn when the trees became dormant. In young trees, we detected a seasonal interconversion between the free and esterified forms of beta-sitosterol and campesterol, and within the steryl ester fraction between squalene and betulaprenol-7. The concentration of esterified sterols/isoprenoids was exceptionally high, especially in the inner regions of mature stem wood (mean 0.6 +/- 0.03% of wood dry mass). No heartwood formation was detected.     

Water relations and gas exchange of Acer saccharum seedlings in contrasting natural light and water regimes

Field measurements were made of leaf photosynthesis (A), stomatal conductance (g) and leaf water relations for sugar maple (Acer saccharum Marsh.) seedlings growing in a forest understory, small gap or large clearing habitat in southwestern Wisconsin, USA. Predawn water status, leaf gas exchange and plasticity in field and laboratory water relations characteristics were compared among contrasting light environments in a wet year (1987) and a dry year (1988) to evaluate possible interactions between light and water availability in these habitats. Leaf water potentials (Psi(leaf)) at predawn and midday were lower for clearing than gap or understory seedlings. Acclimation of tissue osmotic potentials to light environment was observed among habitats but did not occur within any of the habitats in response to prolonged drought. During a summer drought in 1988, decreases in daily maximum g (g(max)) and maximum A (A(max)) in clearing seedlings were correlated with predawn Psi(leaf), which reached a seasonal minimum of -2.0 MPa. Under well-watered conditions, diurnal fluctuations in Psi(leaf) of up to 2.0 MPa in clearing seedlings occurred along with large midday depressions of A and g. In a wet year, strong stomatal responses to leaf-to-air vapor pressure difference (VPD) in sunny habitats were observed over nine diurnal courses of gas exchange measurements on seedlings in a gap and a clearing. Increasing stomatal limitations to photosynthesis appeared to be responsible for the reduction in A at high VPD for clearing seedlings. In understory seedlings, however, low water-use efficiency and development of leaf water deficits in sunflecks was related to reduced stomatal limitations to photosynthesis relative to seedlings in sunny habitats. Predawn Psi(leaf) and VPD appear to be important factors limiting carbon assimilation in sugar maple seedlings in light-saturating irradiances, primarily through stomatal closure. The overall results are consistent with the idea that sugar maple seedlings exhibit "conservative" water use patterns and have low drought tolerance. Leaf water relations and patterns of water use should be considered in studies of acclimation and species photosynthetic performance in contrasting light environments.     

Nitrogen and phosphorus leaching and uptake by container birch seedlings (Betula pendula Roth) grown in three different fertilizations

Leaching of nitrogen (N) and phosphorus (P) through container peatmedium and N and P uptake by seedlings were determined in commercial productionof birch grown with three different types of fertilization. Half of the amountof nutrients was premixed into the peat medium and the other half was appliedasliquid (=fertigated) in the treatment (PF) commonly used in Finland. In twoother treatments all nutrients, either totally (P-VN) or partly (P-N) inslow-release form, were premixed into the peat. Independently of the treatment,the largest proportion (60 to 80%) of total N leached during May and June.During two growing seasons, the amounts of N leached from PF treatments (9 to36kg ha^–1) were as large as the amounts of Nleachedfrom P-VN and P-N treatments (24 to 46 kg ha^–1).Due to fertigations between container trays, however, the total N load perhectare was greatest in the PF treatments. In the soil water 0.5meter beneaththe container area, the N concentration varied from 10 to 60 mgl^–1. The morphological and chemical properties of theseedlings did not differ greatly between treatments.