Differences in the success of sugar maple and red maple seedlings on acid soils are influenced by nutrient dynamics and light environment

Variation in tolerance to nutrient limitations may contribute to the differential success of sugar maple ( Acer saccharum Marsh.) and red maple ( Acer rubrum L.) on acid soils. The objectives of this study were to examine these relationships as influenced by light environment and test whether sensitivity to nutrient stress is mediated by oxidative stress. First-year sugar maple and red seedlings were grown on forest soil cores contrasting in nutrient availability under high or low light intensity. Foliar nutrition, photosynthesis, growth and antioxidant enzyme activity were assessed. Photosynthesis and growth of sugar maple were significantly lower on nutrient-poor soils and were correlated with leaf nutrient status with Ca and P having the strongest influence. For red maple, only chlorophyll content showed sensitivity to the nutrient-poor soils. High light exacerbated the negative effects of nutrient imbalances on photosynthesis and growth in sugar maple. Antioxidant enzyme activity in sugar maple was highest in seedlings growing on nutrient-poor soils and was inversely correlated with photosynthesis, Ca, P, and Mg concentrations. These results suggest that: (1) sugar maple is more sensitive to nutrient stresses associated with low pH soils than red maple; (2) high light increases sugar maple sensitivity to nutrient stress; (3) the negative effects of nutrient imbalances on sugar maple may be mediated by oxidative stress.     

Below-ground respiratory responses of sugar maple and red maple saplings to atmospheric CO2 enrichment and elevated air temperature

The research described in this paper represents a part of a much broader research project with the general objective of describing the effects of elevated [CO2] and temperature on tree growth, physiological processes, and ecosystem-level processes. The specific objective of this research was to examine the below-ground respiratory responses of sugar maple (Acer saccharum Marsh.) and red maple (Acer rubrum L.) seedlings to elevated atmospheric [CO2] and temperature. Red maple and sugar maple seedlings were planted in the ground in each of 12 open-top chambers and exposed from 1994 through 1997 to ambient air or air enriched with 30 Pa CO2,< in combination with ambient or elevated (+4 °C) air temperatures. Carbon dioxide efflux was measured around the base of the seedlings and from root-exclusion zones at intervals during 1995 and 1996 and early 1997. The CO2 efflux rates averaged 0.4 μmol CO2 m-2 s-1 in the root-exclusion zones and 0.75 μmol CO2 m-2 s-1 around the base of the seedlings. Mineral soil respiration in root-exclusion zones averaged 12% higher in the high temperature treatments than at ambient temperature, but was not affected by CO2 treatments. The fraction of total efflux attributable to root + rhizosphere respiration ranged from 14 to 61% in measurements made around red maple plants, and from 35 to 62% around sugar maple plants. Root respiration rates ranged from 0 to 0.94 μmol CO2 s-1 m-2 of soil surface in red maple and from 0 to 1.02 in sugar maple. In both 1995 and 1996 root respiration rates of red maple were highest in high-CO2 treatments and lowest in high temperature treatments. Specific red maple root respiration rates of excised roots from near the soil surface in 1996 were also highest under CO2 enrichment and lowest in high temperature treatments. In sugar maple the highest rates of CO2 efflux were from around the base of plants exposed to both high temperature and high-CO2, even though specific respiration rates were< lowest for this species under the high temperature and CO2 enrichment regime. In both species, patterns of response to treatments were similar in root respiration and root mass, indicating that the root respiration responses were due in part to differences in root mass. The results underscore the need for separating the processes occurring in the roots from those in the forest floor and mineral soil in order to increase our understanding of the effects of global climate change on carbon sequestration and cycling in the below-ground systems of forests.     

Effect of soil K, Ca and Mg saturation and endomycorrhization on growth and nutrient uptake of sugar maple seedlings

Nutrient imbalances of declining sugar maple (Acer saccharum Marsh.) stands in southeastern Quebec have been associated with high exchangeable Mg levels in soils relative to soil K and Ca. A greenhouse experiment was set up to test the hypothesis that the equilibrium between soil exchangeable K, Ca, and Mg ions influences the growth and nutrient status of sugar maple seedlings. Also tested was whether endomycorrhization can alter nutrient acquisition under various soil exchangeable basic cations ratios. Treatments consisted of seven ratios of soil exchangeable K, Ca, and Mg making up a total base saturation of 58%, and a soil inoculation treatment with the endomycorrhizal fungus Glomus versiforme (control and inoculated), in a complete factorial design. Sugar maple seedlings were grown for 3 months in the treated soils. Plant shoot elongation rate, dry biomass and nutrient concentrations in foliage were influenced by the various ratios of soil cations. The predicted plant biomass and foliar K concentration were highest at a soil Ca saturation of 38%, a soil K saturation of 12%, and a soil Mg saturation of 8%. Potassium concentration in foliage was dependent on the level of Ca and Mg saturation in the soil when soil K saturation was close to 12%. Foliar Ca and Mg levels were more dependent on their corresponding levels in soil than foliar K. Colonization by G. versiforme did not influence seedling growth and macronutrient uptake. The results confirm that growth and nutrition of sugar maple are negatively affected by imbalances in exchangeable basic cations in soils.     

Conifer Presence May Negatively Affect Sugar Maple’s Ability to Migrate into the Boreal Forest Through Reduced Foliar Nutritional Status

The discipline of ecology suffers from a lack of knowledge of non-climatic factors (for example, plant–soil, plant–plant and plant–insect interactions) to predict tree species range shifts under climate change. The next generation of simulation models of forest response to climate change must build upon local observations of species interactions and growth along climatic gradients. We examined whether sugar maple (Acer saccharum) seedlings were disadvantaged with respect to soil nutrient uptake under coniferous canopies, as this species would need to migrate northward into conifer-dominated forests in response to climate change. An experimental design was applied to 3 sites, forming the largest possible latitudinal/climatic gradient for sugar maple in Quebec (Canada) and isolating the effect of conifer presence on its seedling’s nutritional status. We tested whether: (1) both soil and climate and (2) presence of conifers affected foliar nutrient levels of sugar maple seedlings. Climate and soil (through pH) strongly affected nutrient availability for sugar maple seedlings and predicted 63.7% of their foliar nutrient variability. When controlling for site effects, we found a significant negative effect of conifers on foliar Ca and Mg levels of maple seedlings, which can adversely affect their overall health and vigour. When considering projected modifications of the forest environment due to climate change, we suggest that northward migration of sugar maple will be negatively affected by the presence of conifers through reduced foliar nutrition.     

Fertilization effects on fineroot biomass, rhizosphere microbes and respiratory fluxes in hardwood forest soils

Fertilizer-induced reductions in CO(2) flux from soil ((F)CO(2)) in forests have previously been attributed to decreased carbon allocation to roots, and decreased decomposition as a result of nitrogen suppression of fungal activity. Here, we present evidence that decreased microbial respiration in the rhizosphere may also contribute to (F)CO(2) reductions in fertilized forest soils. Fertilization reduced (F)CO(2) by 16-19% in 65-yr-old plantations of northern red oak (Quercus rubra) and sugar maple (Acer saccharum), and in a natural 85-yr-old yellow birch (Betula allegheniensis) stand. In oak plots, fertilization had no effects on fine root biomass but reduced mycorrhizal colonization by 18% and microbial respiration by 43%. In maple plots, fertilization reduced root biomass, mycorrhizal colonization and microbial respiration by 22, 16 and 46%, respectively. In birch plots, fertilization reduced microbial respiration by 36%, but had variable effects on root biomass and mycorrhizal colonization. In plots of all three species, fertilization effects on microbial respiration were greater in rhizosphere than in bulk soil, possibly as a result of decreased rhizosphere carbon flux from these species in fertile soils. Because rhizosphere processes may influence nutrient availability and carbon storage in forest ecosystems, future research is needed to better quantify rhizo-microbial contributions to (F)CO(2).     

Foliar δ15N is affected by foliar nitrogen uptake, soil nitrogen, and mycorrhizae along a nitrogen deposition gradient

Foliar nitrogen isotope (δ¹⁵N) composition patterns have been linked to soil N, mycorrhizal fractionation, and within-plant fractionations. However, few studies have examined the potential importance of the direct foliar uptake of gaseous reactive N on foliar δ¹⁵N. Using an experimental set-up in which the rate of mycorrhizal infection was reduced using a fungicide, we examined the influence of mycorrhizae on foliar δ¹⁵N in potted red maple (Acer rubrum) seedlings along a regional N deposition gradient in New York State. Mycorrhizal associations altered foliar δ¹⁵N values in red maple seedlings from 0.06 to 0.74 ‰ across sites. At the same sites, we explored the predictive roles of direct foliar N uptake, soil δ¹⁵N, and mycorrhizae on foliar δ¹⁵N in adult stands of A. rubrum, American beech (Fagus grandifolia), black birch (Betula lenta), and red oak (Quercus rubra). Multiple regression analysis indicated that ambient atmospheric nitrogen dioxide (NO₂) concentration explained 0, 69, 23, and 45 % of the variation in foliar δ¹⁵N in American beech, red maple, red oak, and black birch, respectively, after accounting for the influence of soil δ¹⁵N. There was no correlation between foliar δ¹³C and foliar %N with increasing atmospheric NO₂ concentration in most species. Our findings suggest that total canopy uptake, and likely direct foliar N uptake, of pollution-derived atmospheric N deposition may significantly impact foliar δ¹⁵N in several dominant species occurring in temperate forest ecosystems.     

Effects of elevated ozone and low light on diurnal and seasonal carbon gain in sugar maple

The long‐term interactive effects of ozone and light on whole‐tree carbon balance of sugar maple (Acer saccharum Marsh.) seedlings were examined, with an emphasis on carbon acquisition, foliar partitioning into starch and soluble sugars, and allocation to growth. Sugar maple seedlings were fumigated with ambient, 1·7 × ambient and 3·0 × ambient ozone in open‐top chambers for 3 years under low and high light (15 and 35% full sunlight, respectively). Three years of ozone fumigation reduced the total biomass of seedlings in the low‐ and high‐light treatments by 64 and 41%, respectively, but had no effect on whole‐plant biomass allocation. Ozone had no effect on net photosynthesis until late in the growing season, with low‐light seedlings generally exhibiting more pronounced reductions in photosynthesis. The late‐season reduction in photosynthesis was not due to impaired stomatal function, but was associated more with accelerated senescence or senescence‐like injury. In contrast, the 3·0 × ambient ozone treatment immediately reduced diurnal starch accumulation in leaves by over 50% and increased partitioning of total non‐structural carbohydrates into soluble sugars, suggesting that injury repair processes may be maintaining photosynthesis in late spring and early summer at the expense of storage carbon. The results in the present study indicate that changes in leaf‐level photosynthesis may not accurately predict the growth response of sugar maple to ozone in different light environments. The larger reduction in seedling growth under low‐light conditions suggests that seedlings in gap or closed‐canopy environments are more susceptible to ozone than those in a clearing. Similarly, understanding the effects of tropospheric ozone on net carbon gain of a mature tree will require scaling of leaf‐level responses to heterogeneous light environments, where some leaves may be more susceptible than others.     

Foliar and wood chemistry of sugar maple along a gradient of soil acidity and stand health

The decline of sugar maple (Acer saccharum Marsh.) in forest of north-eastern North America is an important environmental issue. In this study, relationships between, soil, wood and foliar chemistry were assessed for 17 stands distributed within a large area of the Quebec sugar maple forest and that were growing on soils with a strong gradient of acidity and base saturation. There were many significant relationships between variables describing the acid-base status of the top-B soil (Ca and Mg concentrations, exchangeable acidity and base saturation) and Ca and Mn concentrations and Ca/Mn and Mg/Mn in tree tissues. Manganese was the element that showed the strongest inverse non-linear relationships with top-B soil base saturation with variance explanation of 71 and 65%, for wood and foliage, respectively. The 17 sites were divided in two groups according to their level of decline. The declining stands had significantly higher wood Mn and Mg concentrations and lower Ca/Mn ratios and significantly higher foliar Mn and lower Ca and Al concentrations. It was impossible to determine if these differences were a cause or a symptom of sugar maple health. However, the increase in Mn concentrations in tree tissues with increasing soil acidity, as well as the higher Mn concentrations in declining as compared to healthy stands suggest that Mn, as well as low Ca availability, could be an important contributing factor in the sugar maple decline.     

Influence of experimental snow removal on root and canopy physiology of sugar maple trees in a northern hardwood forest

Due to projected increases in winter air temperatures in the northeastern USA over the next 100 years, the snowpack is expected to decrease in depth and duration, thereby increasing soil exposure to freezing air temperatures. To evaluate the potential physiological responses of sugar maple (Acer saccharum Marsh.) to a reduced snowpack, we measured root injury, foliar cation and carbohydrate concentrations, woody shoot carbohydrate levels, and terminal woody shoot lengths of trees in a snow manipulation experiment in New Hampshire, USA. Snow was removed from treatment plots for the first 6 weeks of winter for two consecutive years, resulting in lower soil temperatures to a depth of 50 cm for both winters compared to reference plots with an undisturbed snowpack. Visibly uninjured roots from trees in the snow removal plots had significantly higher (but sub-lethal) levels of relative electrolyte leakage than trees in the reference plots. Foliar calcium: aluminum (Al) molar ratios were significantly lower, and Al concentrations were significantly higher, in trees from snow removal plots than trees from reference plots. Snow removal also reduced terminal shoot growth and increased foliar starch concentrations. Our results are consistent with previous research implicating soil freezing as a cause of soil acidification that leads to soil cation imbalances, but are the first to show that this translates into altered foliar cation pools, and changes in soluble and structural carbon pools in trees. Increased soil freezing due to a reduced snowpack could exacerbate soil cation imbalances already caused by acidic deposition, and have widespread implications for forest health in the northeastern USA.     

Decline of arbuscular mycorrhizal fungi in northern hardwood forests exposed to chronic nitrogen additions

Arbuscular mycorrhizal (AM) fungi are important below-ground carbon (C) sinks that can be sensitive to increased nitrogen (N) availability. The abundance of AM fungi (AMF) was estimated in maple (Acer spp.) fine roots following more than a decade of experimental additions of N designed to simulate chronic atmospheric N deposition. Abundance of AMF was measured by staining and ocular estimation, as well as by analyzing for the AMF indicator fatty acid 16:1omega5c in phospholipid (biomass indicator) and neutral lipid (lipid storage indicator) fractions. Arbuscular mycorrhizal fungal biomass, storage structures and lipid storage declined in response to N addition measured by both methods. This pattern was found when AM response was characterized as colonization intensity, on an areal basis and in proportion to maple above-ground biomass. The phospholipid fraction of the fatty acid 16:1omega5c was positively correlated with total AMF colonization and the neutral lipid fraction with vesicle colonization. Decreased AMF abundance with simulated N deposition suggests reduced C allocation to these fungi or a direct soil N-mediated decline. The fatty acid (phospholipid and neutral lipid fractions) 16:1omega5c was found to be a good indicator for AMF active biomass and stored energy, respectively.     

Evidence for oxidative stress in sugar maple stands growing on acidic,nutrient imbalanced forest soils

Soil acidification and the disruption of nutrient cycles appear to be important factors that weaken sugar maple resistance to both abiotic and biotic stresses and predispose it to decline symptoms. Although connections between edaphic stress and decline symptoms have been identified, very little is known about the physiological and biochemical mechanisms that underlie this relationship. In this study, we tested the hypothesis that foliar nutrient imbalances impair the photosynthetic apparatus of sugar maple through oxidative stress. We examined leaf nutrition, photosynthesis and antioxidant enzyme activity (a biomarker of oxidative stress) from early June to late August in three-paired overstory sugar maple stands on Pennsylvania’s Allegheny Plateau that contrast in soil nutrient availability according to slope position. Beginning in early June, trees on upper slopes (nutrient-poor) had significantly lower foliar Ca and Mg concentrations and significantly higher foliar Mn concentrations than trees on lower slopes. These differences increased throughout summer peaking in late August. Photosynthesis and antioxidant enzyme activity closely reflected changes in foliar nutrient status throughout the summer. In the latter half of the summer, leaf gas exchange and chlorophyll content were significantly lower and antioxidant enzyme activity was significantly higher in stands on upper slope soils. At the end of August, leaf nutrient imbalances corresponded with lower rates of photosynthesis and higher antioxidant enzyme activity, suggesting that foliar nutrient imbalances may impair sugar maple function through mechanisms of oxidative stress.     

The effects of acid deposition on the biogeochemical cycles of major nutrients in miniature red spruce ecosystems

As part of an experimental study of air pollution effects on tree growth and health, we combined process studies with an ecosystem approach to evaluate the effects of acidic deposition on soil acidification, nutrient cycling and proton fluxes in miniature red spruce ecosystems. Ninety red spruce saplings were transplanted into 1-m diameter pots containing reconstructed soil profiles and exposed to simulated acid rain treatments of pH 3.1, 4.1 and 5.1 for four consecutive growing seasons. All the principal fluxes of the major elements were measured. During the first year of treatments, the disturbance associated with the transplanting of the experimental trees masked any treatment effects by stimulating N mineralization rates and consequent high N0₃ ⁻ cation, and H⁺ flux through the soil profile. In subsequent years, leaching of base cations and labile Al was accelerated in the most intensive acid treatment and corresponding declines in soil pH and exchangeable pools of Ca and Mg and increases in exchangeable Al concentrations were observed in the organic horizon. Leaching of Ca²⁺ and Mg²⁺ also was significantly higher in the pH 4.1 than in the pH 5.1 treatment. Flux of Ca from foliage and soil was increased in response to strong acid loading and root uptake increased to compensate for foliar Ca losses. In contrast, K cycling was dominated by root uptake and internal cycling and was relatively insensitive to strong acid inputs. Cation leaching induced by acidic deposition was responsible for the majority of H⁺ flux in the pH 3.1 treatment in the organic soil horizon whereas root uptake accounted for most of the H⁺ flux in the pH 4.1 and 5.1 treatments. Although no measurable effects on tree nutrition or health were observed, base cation leaching was significantly accelerated by acidic deposition, even at levels below that observed in the eastern U.S., warranting continued concern about acid deposition effects on the soil base status of forested ecosystems.     

Testing the enemy release hypothesis: a comparison of foliar insect herbivory of the exotic Norway maple (Acer platanoides L.) and the native sugar maple (A. saccharum L.)

Norway maple (Acer platanoides) is a Eurasian introduced tree species which has invaded the North American range of its native congener, sugar maple (A. saccharum). One hypothesis used to explain the success of an invasive species is the enemy release hypothesis (ERH), which states that invasive species are often particularly successful in their new range because they lack the enemies of their native range. In this study, we hypothesized that Norway maple would have less insect damage than sugar maple due to such enemy release. Autumn 2005 and summer 2006 leaves of Norway and sugar maple were collected from six sites in New Jersey and Pennsylvania to compare percent leaf area loss, gall damage, fungal damage, and specific leaf area (cm²/g). Although both species had low overall mean levels of leaf damage (0.4–2.5%), in both years/seasons Norway maple had significantly less leaf damage than sugar maple. Insects were also collected to compare insect assemblies present on each tree species. The numbers of insect taxa and individuals found on each species were nearly equivalent. Overall, the results of this study are consistent with the enemy release hypothesis for Norway maple. In addition, sugar maples when surrounded by Norway maples tended to show reduced herbivory. This suggests that the spread of Norway maple in North America, by reducing amounts of insect herbivory, may have further ecosystem-wide impacts.     

Effect of snow removal on leaf water potential,soil moisture,leaf and soil nutrient status and leaf peroxidase activity of sugar maple

Effect of removal of snow cover in winter was investigated in an 80-year-old sugar maple (Acer saccharum Marsh.) stand in southern Quebec. We hypothesized that winter soil frost would induce some of the decline symptoms observed in sugar maple stands in southern Quebec in the early 1980's. Snow was continuously removed from around trees for a one week (partial removal) or for a four-month period (complete removal) during the 1990–1991 winter. Foliage and soils were sampled periodically during the summer of 1991. The complete snow removal treated trees showed decreased leaf water potential and increased peroxidase activity over most of the growing season. Foliar Ca was reduced in both snow removal treatments early in the growing season while foliar N was reduced in the complete snow removal trees late in the growing season. Soil NO ₃ ^– and K⁺ were elevated in both snow removal treatments at various times throughout the growing season. Prolonged soil frost in a sugar maple stand can induce lower leaf water potential, higher leaf peroxidase activity and early leaf senescence during the following growing season. Soil frost may have reduced nutrient uptake without affecting significantly the leaf nutrient status.     

Effects of base cation fertilization on the nutrient status,free amino acids and some carbon fractions of the leaves of sugar maple (Acer saccharum Marsh.)

A study was conducted in the Lower Laurentians of southern Quebec to test the hypothesis that base cation fertilization would improve the nutrient status of declining sugar maples (Acer saccharum Marsh.) and alter the partitioning of leaf C and N. Six 40×40 m plots were delineated in an 80 year old stand of sugar maple. Three plots received a mixture of fertilizer and liming materials (500 kg ha^–1 of K₂SO₄, 250 kg ha^–1 of CaCO₃ and 250 kg ha^–1 of CaMg(CO₂)₂) in the spring of 1989. Leaves from mid crown of dominant or co-dominant maples were sampled monthly during the 1990 growing season. Trees were cored in 1992 to measure their response in diameter growth. Fertilization increased diameter growth and foliar K concentration of trees but reduced foliar Ca concentration. Fertilization resulted in lower starch concentrations and higher ratios of soluble sugars to starch in June and September, and in higher free amino acid concentrations but lower ratio of total non-structural carbohydrates to free amino acids in September. Leaf proline concentration was correlated with leaf starch concentration (r=0.39). The results suggest that amelioration of K deficiencies in sugar maple through fertilization with a mixture of base cations can increase tree growth and affect the seasonal dynamics of foliar C and N pools.Abbreviations FAA free amino acids - TNC total non-structural carbohydrates     

Base-cation Cycling by Individual Tree Species in Old-growth Forests of Upper Michigan,USA

The influence of individual tree species on base-cation (Ca, Mg, K, Na) distribution and cycling was examined in sugar maple (Acer saccharum Marsh.), basswood (Tilia americana L.), and hemlock (Tsuga canadensis L.) in old-growth northern hardwood – hemlock forests on a sandy, mixed, frigid, Typic Haplorthod over two growing seasons in northwestern Michigan. Base cations in biomass, forest floor, and mineral soil (0–15 cm and 15–40 cm) pools were estimated for five replicated trees of each species; measured fluxes included bulk precipitation, throughfall, stemflow, litterfall, forest-floor leachate, mineralization + weathering, shallow-soil leachate, and deep-soil leachate. The three species differed in where base cations had accumulated within the single-tree ecosystems. Within these three single-tree ecosystems, the greatest quantity of base cations in woody biomass was found in sugar maple, whereas hemlock and basswood displayed the greatest amount in the upper 40 cm of mineral soil. Base-cation pools were ranked: sugar maple > basswood, hemlock in woody biomass; sugar maple, basswood > hemlock in foliage; hemlock > sugar maple, basswood in the forest floor, and basswood > sugar maple, hemlock in the mineral soil. Base-cation fluxes in throughfall, stemflow, the forest-floor leachate, and the deep-soil leachate (2000 only) were ranked: basswood > sugar maple > hemlock. Our measurements suggest that species-related differences in nutrient cycling are sufficient to produce significant differences in base-cation contents of the soil over short time intervals (<65 years). Moreover, these species-mediated differences may be important controls over the spatial pattern and edaphic processes of northern hardwood-hemlock ecosystems in the upper Great Lakes region.     

Fine root chemistry and decomposition in model communities of north-temperate tree species show little response to elevated atmospheric CO<Subscript>2</Subscript> and varying soil resource availability

Rising atmospheric [CO₂] has the potential to alter soil carbon (C) cycling by increasing the content of recalcitrant constituents in plant litter, thereby decreasing rates of decomposition. Because fine root turnover constitutes a large fraction of annual NPP, changes in fine root decomposition are especially important. These responses will likely be affected by soil resource availability and the life history characteristics of the dominant tree species. We evaluated the effects of elevated atmospheric [CO₂] and soil resource availability on the production and chemistry, mycorrhizal colonization, and decomposition of fine roots in an early- and late-successional tree species that are economically and ecologically important in north temperate forests. Open-top chambers were used to expose young trembling aspen (Populus tremuloides) and sugar maple (Acer saccharum) trees to ambient (36 Pa) and elevated (56 Pa) atmospheric CO₂. Soil resource availability was composed of two treatments that bracketed the range found in the Upper Lake States, USA. After 2.5 years of growth, sugar maple had greater fine root standing crop due to relatively greater allocation to fine roots (30% of total root biomass) relative to aspen (7% total root biomass). Relative to the low soil resources treatment, aspen fine root biomass increased 76% with increased soil resource availability, but only under elevated [CO₂]. Sugar maple fine root biomass increased 26% with increased soil resource availability (relative to the low soil resources treatment), and showed little response to elevated [CO₂]. Concentrations of N and soluble phenolics, and C/N ratio in roots were similar for the two species, but aspen had slightly higher lignin and lower condensed tannins contents compared to sugar maple. As predicted by source-sink models of carbon allocation, pooled constituents (C/N ratio, soluble phenolics) increased in response to increased relative carbon availability (elevated [CO₂]/low soil resource availability), however, biosynthetically distinct compounds (lignin, starch, condensed tannins) did not always respond as predicted. We found that mycorrhizal colonization of fine roots was not strongly affected by atmospheric [CO₂] or soil resource availability, as indicated by root ergosterol contents. Overall, absolute changes in root chemical composition in response to increases in C and soil resource availability were small and had no effect on soil fungal biomass or specific rates of fine root decomposition. We conclude that root contributions to soil carbon cycling will mainly be influenced by fine root production and turnover responses to rising atmospheric [CO₂], rather than changes in substrate chemistry.     

The effects of liming and gypsum applications on a sessile oak (Quercus petraea (M.) Liebl.) stand at La Croix-Scaille (French Ardennes) I. Site characteristics, soil chemistry and aerial biomass

In a former 45 to 50 year old sessile oak ( Quercus petraea (M.) Liebl.) coppice mixed with birch (Betula pubescens Ehrh.) and rowan (Sorbus aucuparia L.) on a poor acidic forest soil at la Croix-Scaille in the French Ardennes, several liming amendments were applied in 1990 and 1994. Data on soil and soil solution composition, as well as stand growth and foliar composition were collected between 1994 and 1997. All treatments, containing 1.4 t ha-1 equivalent of CaO supplied as lime, gypsum or a mixture of the two, resulted in an increase of cation exchange capacity and base saturation down to 15 cm and for CaSO4 treatments down to 30 to 45 cm, increases of soil pH and Ca concentration at the surface and a decrease of Al concentration in the soil and soil solution in the surface layers. No negative effects like increased nitrate or cation leaching were observed. Although Mg nutrition was not improved by the treatments (not containing Mg), a relative and maintained gain of radial increment of sessile oak in the order of 40% for both lime and gypsum applied, was observed immediately from the first year on, after the application (1991).     

Soil Base Saturation Combines with Beech Bark Disease to Influence Composition and Structure of Sugar Maple-Beech Forests in an Acid Rain-Impacted Region

Sugar maple, an abundant and highly valued tree species in eastern North America, has experienced decline from soil calcium (Ca) depletion by acidic deposition, while beech, which often coexists with sugar maple, has been afflicted with beech bark disease (BBD) over the same period. To investigate how variations in soil base saturation combine with effects of BBD in influencing stand composition and structure, measurements of soils, canopy, subcanopy, and seedlings were taken in 21 watersheds in the Adirondack region of NY (USA), where sugar maple and beech were the predominant canopy species and base saturation of the upper B horizon ranged from 4.4 to 67%. The base saturation value corresponding to the threshold for Al mobilization (16.8%) helped to define the species composition of canopy trees and seedlings. Canopy vigor and diameter at breast height (DBH) were positively correlated (P < 0.05) with base saturation for sugar maple, but unrelated for beech. However, beech occupied lower canopy positions than sugar maple, and as base saturation increased, the average canopy position of beech decreased relative to sugar maple (P < 0.10). In low-base saturation soils, soil-Ca depletion and BBD may have created opportunities for gap-exploiting species such as red maple and black cherry, whereas in high-base saturation soils, sugar maple dominated the canopy. Where soils were beginning to recover from acidic deposition effects, sugar maple DBH and basal area increased progressively from 2000 to 2015, whereas for beech, average DBH did not change and basal area did not increase after 2010.     

Norway Maple (Acer platanoides) Invasion of a Natural Forest Stand: Understory Consequence and Regeneration Pattern

Norway maple (Acer platanoidesis) is invasive in a natural stand in suburban Ithaca, NY. To determine the understory pattern and consequences of a Norway maple invasion, I compared density and species richness under Norway maples and sugar maples (Acer saccharum). Mean sapling density was significantly lower (P<0.0027) under Norway maples (3.64/100 m²±1.6 SE) than under sugar maples (19.4/100 m²±4.4 SE). Mean sapling species richness was significantly lower (P<0.0018) under Norway maples (0.7/32 m²±0.18 SE) than under sugar maples (2.6/32 m²±0.48 SE). Likewise, Norway maple regeneration is more frequent under sugar maples than sugar maple regeneration: 57% of sugar maple plots had Norway maple saplings while 0% of Norway maple plots had sugar maple saplings. Two significant plot effects were found for presence–absence: Norway maple saplings grow under Norway maples with a significantly lower frequency (P<0.03) than under sugar maples; sugar maple saplings grow under Norway maples with a significantly lower frequency (P<0.000) than under sugar maples. Across the site, Norway maple saplings were the most abundant (29 saplings for 480 m²). The success of Norway maple regeneration and the reductions in total stem density beneath Norway maples is most likely the result of its strong competitive abilities, notably its high shade tolerance and abundant seed crops.