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.     

Use of foliar Ca/Sr discrimination and <Superscript>87</Superscript>Sr/<Superscript>86</Superscript>Sr ratios to determine soil Ca sources to sugar maple foliage in a northern hardwood forest

Calcium/strontium and ⁸⁷Sr/⁸⁶Sr ratios in foliage can be used to determine the relative importance of different soil sources of Ca to vegetation, if the discrimination of Ca/Sr by the plant between nutrient sources and foliage is known. We compared these tracers in the foliage of sugar maple (Acer saccharum) to the exchange fraction and acid leaches of soil horizons at six study sites in the White Mountains of New Hampshire, USA. In a previous study, sugar maple was shown to discriminate for Ca compared to Sr in foliage formation by a factor of 1.14 ± 0.12. After accounting for the predicted 14% shift in Ca/Sr, foliar Ca/Sr and ⁸⁷Sr/⁸⁶Sr ratios closely match the values in the Oie horizon at each study site across a 3.6-fold variation in foliar Ca/Sr ratios. Newly weathered cations, for which the Ca/Sr and ⁸⁷Sr/⁸⁶Sr ratios are estimated from acid leaches of soils, can be ruled out as a major Ca source to current foliage. Within sites, the ⁸⁷Sr/⁸⁶Sr ratio of the soil exchange pool in the Oa horizon and in the 0–10 cm and 10–20 cm increments of the mineral soil are similar to the Oie horizon and sugar maple foliar values, suggesting a common source of Sr in all of the actively cycling pools, but providing no help in distinguishing among them as sources to foliage. The Ca/Sr ratio in the soil exchange pool, however, decreases significantly with depth, and based on this variation, the exchange pool below the forest floor can be excluded as a major Ca source to the current sugar maple foliage. This study confirms that internal recycling of Ca between litter, organic soil horizons and vegetation dominate annual uptake of Ca in northern hardwood ecosystems. Refinement of our understanding of Ca and Sr uptake and allocation in trees allows improvement in the use of Ca/Sr and ⁸⁷Sr/⁸⁶Sr ratios to trace Ca sources to plants.     

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.     

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.     

Site factors,foliar nutrient levels and growth of Cordia alliodora plantations in the humid lowlands of Northern Costa Rica

Within the perhumid, Atlantic lowlands of northern Costa Rica, Cordia alliodora plantations were studied in order to explain the observed pattern of growth irregularities. The soils, that were partly used as pastures over long periods, could be classified roughly into two units: (i) red, deeply weathered, slightly acidic soils from Mg-enriched parent material and (ii) brown, strongly acidic soils with high saturation of exchangeable Al (up to 80%).Leaf analysis revealed that Cordia is a highly demanding species in respect to macronutrients. Poorly growing trees in slope position suffered from an insufficient supply of N and P. K/Mg ratios of chlorotic leaves are very low. Soil analysis showed that nutrient deficiencies were related to (i) insufficient soil nutrient reserves of the poor, tropical soils or (ii) to an inhibition of nutrient uptake by soil physical or chemical factors.All sites are characterized by very low K reserves and losses of nutrients that are organically bound (N, P) caused by erosion. In the Mg-rich red soils, low amounts of K lead to K/Mg imbalances. Soil compaction caused by cattle grazing occurs on both soil units. In negatively influences the root development of Cordia, and hence nutrient uptake. In the brown soils, high amounts of exchangeable Al hinder a sufficient supply of nutrients (e.g.P) to the assimilation organs.     

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.     

Base cation stimulation of mycorrhization and photosynthesis of sugar maple on acid soils are coupled by foliar nutrient dynamics

The nutritional benefits that mycorrhizal associations provide to plants may be constrained by acidic soil conditions resulting in decreased photosynthetic function. Sugar maple (Acer saccharum) and red maple (Acer rubrum) seedlings were grown on a native acidic (pH 4.1) soil both unamended and amended with base cations (pH 6.2). In a second study a fungicide treatment was included. Foliar nutrition, mycorrhizal colonization, photosynthesis and their relationships were assessed. On the native soil, red maple maintained higher levels of mycorrhizal colonization and photosynthesis than sugar maple but showed little response to base cation amendments. Mycorrhizal colonization and photosynthesis of sugar maple increased significantly in response to base cation amendments. Correlations were observed among mycorrhizal colonization, foliar nutrition and photosynthesis. The fungicide treatment indicated that 50% of the base cation-induced increase in sugar maple photosynthesis was mycorrhiza related. The results suggest that base cation stimulation of mycorrhization and photosynthesis of sugar maple on acid soils are coupled by foliar nutrient dynamics. Red maple exhibits much less sensitivity to these same edaphic conditions.     

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.     

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.     

Root biomass distribution in a moist tropical montane forest

Effects of N sources (ammonium, nitrate and ammonitrate) and VA mycorrhizae (Glomus intraradices) on rhizosphere soil characteristics (pH, exchangeable acidity, exchangeable cations, inorganic N concentrations) growth and nutrient acquisition of coffee seedlings (Coffea arabica L. cv guatemala) were investigated in a pot study with an acid soil (Red Bluff Loam) sterilized by autoclaving. Ammonium addition decreased rhizosphere pH while nitrate and ammonitrate additions both increased rhizosphere pH. Mycorrhizae induced a higher pH, a lower exchangeable acidity and higher values of exchangeable cations in the rhizosphere. Ammonium addition resulted in a lower mycorrhizal infection than the two other N sources. Mycorrhizal plants grew better and accumulated more N, Ca and Mg than non-mycorrhizal plants.     

祁连山西水林区土壤阳离子交换量及盐基离子的剖面分布

以祁连山西水林区分布的棕钙土、灰褐土、栗钙土和高山草甸土为对象,研究了阳离子交换量和盐基离子(K+、Na+、Ca2+、Mg2+)的剖面分布规律及其与土壤理化因子的关系。结果表明:土壤阳离子交换量(CEC,介于4.80—48.10 cmol/kg)和盐基总量(TEB,介于4.67—21.34 cmol/kg)随剖面深度的增加逐渐减小,不同土壤类型的大小顺序为:灰褐土>高山草甸土>栗钙土>棕钙土;土壤盐基组成以Ca2+、Mg2+为主(占TEB的比例平均为71.6%、22.9%),K+、Na+所占比例较低(占TEB的比例平均为3.3%、2.2%);棕钙土、灰褐土和栗钙土盐基离子的剖面分布由浅至深呈现:K+≈Ca2+>Na+≈Mg2+,高山草甸土盐基离子则呈现:K+>Na+>Mg2+>Ca2+。不同土壤类型间盐基离子的含量及饱和度随发生层次不同存在较大差异。土壤有机质是CEC的主要贡献因素,粉粒对CEC也有显著的促进作用,而砂粒、CaCO3对CEC有显著抑制作用。土壤生物复盐基作用弱于淋溶作用,造成盐基饱和度较大(BSP,介于44.4%—97.2%),并随剖面深度的增加逐渐增大。相关性分析表明,土壤交换性Na+、Mg2+的含量及饱和度均呈极显著正相关,交换性Na+、Mg2+饱和度与CaCO3含量呈极显著正相关;pH值与BSP呈极显著正相关;土壤速效P含量与CEC呈极显著正相关,速效K含量与交换性K+含量呈极显著正相关。     

Coupled Nitrogen and Calcium Cycles in Forests of the Oregon Coast Range

Nitrogen (N) is a critical limiting nutrient that regulates plant productivity and the cycling of other essential elements in forests. We measured foliar and soil nutrients in 22 young Douglas-fir stands in the Oregon Coast Range to examine patterns of nutrient availability across a gradient of N-poor to N-rich soils. N in surface mineral soil ranged from 0.15 to 1.05% N, and was positively related to a doubling of foliar N across sites. Foliar N in half of the sites exceeded 1.4% N, which is considered above the threshold of N-limitation in coastal Oregon Douglas-fir. Available nitrate increased five-fold across this gradient, whereas exchangeable magnesium (Mg) and calcium (Ca) in soils declined, suggesting that nitrate leaching influences base cation availability more than soil parent material across our sites. Natural abundance strontium isotopes (⁸⁷Sr/⁸⁶Sr) of a single site indicated that 97% of available base cations can originate from atmospheric inputs of marine aerosols, with negligible contributions from weathering. Low annual inputs of Ca relative to Douglas-fir growth requirements may explain why foliar Ca concentrations are highly sensitive to variations in soil Ca across our sites. Natural abundance calcium isotopes (δ⁴⁴Ca) in exchangeable and acid leachable pools of surface soil measured at a single site showed 1 per mil depletion relative to deep soil, suggesting strong Ca recycling to meet tree demands. Overall, the biogeochemical response of these Douglas-fir forests to gradients in soil N is similar to changes associated with chronic N deposition in more polluted temperate regions, and raises the possibility that Ca may be deficient on excessively N-rich sites. We conclude that wide gradients in soil N can drive non-linear changes in base-cation biogeochemistry, particularly as forests cross a threshold from N-limitation to N-saturation. The most acute changes may occur in forests where base cations are derived principally from atmospheric inputs.     

Influence of calcium, potassium, and magnesium on Cornus florida L. density and resistance to dogwood anthracnose

Dogwood anthracnose, a major disease of Cornus florida L., has caused heavy mortality of C. florida in eastern United States forests. Disease severity and rate of infection have been shown to vary with several environmental factors, but the link between soil cation availability and anthracnose has not been examined. We hypothesized that soil cation availability, particularly calcium (Ca), potassium (K), and magnesium (Mg), would influence dogwood survival from anthracnose. In forested stands, positive correlations between soil Ca, K, and Mg saturation and C. florida stem density and basal area were found. The effect of these cations at four levels (0, 50, 100, and 200%) of a standard nursery fertilization rate on C. florida seedling survival and resistance to dogwood anthracnose was tested. Although most of the seedlings died after one season of exposure to dogwood anthracnose, seedlings that had lower inputs of Ca and K cations showed higher levels of disease severity sooner than seedlings in other treatments, suggesting that these nutrients play a role in C. florida survival from anthracnose. Magnesium treatment levels did not appear to have an effect on C. florida disease severity or mortality.     

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.     

Influence of Norway spruce seedlings on the nutrient availability in mineral soil and forest floor material

A greenhouse experiment was performed to evaluate the effect of Norway spruce (Picea abies (L.) Karst.) seedlings on net nutrient availability in five different growing media containing F- or H-layer and mineral soil originating from a haplic podzol in northern Sweden. The initial total amounts of eight nutrient elements (N, K, P, Ca, Mg, Mn, Fe, Zn) and exchangeable amounts of the same elements were analyzed in pots with or without spruce seedlings. In the planted pots seedling nutrient uptake was also estimated. After 26 weeks, higher net nutrient availability with seedlings was found in 25 out of the 40 (62%) growing media and nutrient element combinations. A positive seedling effect on net nutrient availability might be explained by rhizodeposition stimulating the soil microorganism activity and accelerating the weathering of minerals or by seedling roots promoting the nutrient providing processes through changes in soil chemical and physical properties. Nitrogen availability was primarily affected by what part of the forest floor the growing medium contained although the positive response to seedling presence was apparent. The positive net availability response of P, Ca, Mg, Mn, Fe and Zn to seedling presence was on the other hand relatively strong. In the case of P, K, and Zn the growing medium composition (if the F- and H-layer was pure or mixed with mineral soil) was also an important factor for the estimated net availability. Pure F-and H-layer provided greater P- and K-availability while the availability of Zn increased when mineral soil was added. The influence of growing plants ought to be considered when soil samples are used for assessing the nutrient availability.     

Tree Species Effects on Calcium Cycling: The Role of Calcium Uptake in Deep Soils

Soil acidity and calcium (Ca) availability in the surface soil differ substantially beneath sugar maple (Acer saccharum) and eastern hemlock (Tsuga canadensis) trees in a mixed forest in northwestern Connecticut. We determined the effect of pumping of Ca from deep soil (rooting zone below 20-cm mineral soil) to explain the higher available Ca content in the surface soil beneath sugar maple. We measured the atmospheric input of Ca with bulk deposition collectors and estimated Ca weathering and Ca mineralization in the surface soil (rooting zone above 20-cm mineral soil) from strontium isotope measurements and observed changes in exchangeable Ca in soils during field incubation. Calcium leaching at 20 cm was calculated by combining modeled hydrology with measured Ca soil solution concentrations at 20-cm depth. We measured root length distribution with depth beneath both tree species. Calcium leaching from the surface soil was much higher beneath sugar maple than hemlock and was positively related with the amount of Ca available in the surface soil. Calcium leaching from the surface soil beneath sugar maple was higher than the combined Ca input from atmospheric deposition and soil weathering. Without Ca uptake in the deep soil, surface soils are being depleted in Ca, especially beneath sugar maple. More organically bound Ca was mineralized beneath sugar maple than beneath hemlock. A relatively small part of this Ca release was leached below the surface soil, suggesting that, beneath both tree species, most of the Ca cycling is occurring in the surface soil. Sugar maple had more fine roots in the deep soil than hemlock and a greater potential to absorb Ca in the deep soil. With a simple model, we showed that a relatively small amount of Ca uptake in the deep soil beneath sugar maple is able to sustain high amounts of available Ca in the surface soil. Received 20 June 2001; accepted 6 December 2001.     

Influence of excess nitrogen deposition on a white spruce (Picea glauca) stand in southern Alaska

Excess N delivered to forest ecosystems has been shown to alterinternal ecosystem biogeochemical cycles, contribute to forestdecline, and negatively affect the health of receiving waters.In the vicinity of the Nikiski Industrial Complex, Kenai Peninsula,Alaska, there has been recent concern about the influence ofNH3 emissions that have occurred for over twodecades on local soils and vegetation. The study site representedan opportunity to examine the influence of elevated N depositionon a northern coniferous ecosystem in an area with a low backgroundof N deposition. Overstory vegetation in the area is dominated bywhite spruce (Picea glauca Moench. Voss) and paper birch (Betulapapyrifera Marsh.). Mortality of both species has occurred adjacent(<2 km) to the industrial complex. Average annual Ndeposition rates ranged from 0.7 to 21.0 kg ha-1 y-1in the area, with the highest rates closest to the complex. Sulfatedeposition at the site was low. Due to the high NH3deposition, precipitation near the complex was less acidic thanprecipitation in general; bulk precipitation pH ranged from 5.51to 7.06. Within 1.80 km of the facility there was an increase inKCl- and resin- extractable soil NH4+ andNO3- in the O horizon, and a decrease in soil pHcompared to soils further from the facility. Spruce near thefacility had chlorotic foliage and thinning crowns, higherconcentrations of N, but lower foliar Ca and Mg. Foliar Mglevels approached deficiency levels, but foliar Ca was wellabove reported deficiency levels at all sites. Both Mg:N andCa:N ratios, however, suggest nutrient imbalances in the highN deposition zone. Canopy death and fertilization by N appearto have encouraged growth of the native bluejoint grass. Thepresence of elevated NO3- in O horizon soilextracts, elevated NO3- in resin bags placed betweenthe O and E horizons, and nutritional imbalances in the foliagesuggest that N saturation may be occurring in soils adjacent to thefacility.     

Wood-ash recycling affects forest soil and tree fine-root chemistry and reverses soil acidification

Wood ash was applied to a forest ecosystem with the aim to recycle nutrients taken from the forest and to mitigate the negative effects of intensive harvesting. After two years, the application of 8,000 kg ha⁻¹ of wood ash increased soil exchangeable Ca and Mg. Similarly, an increase in Ca and Mg in the Norway spruce fine roots was recorded, leading to significant linear correlations between soil and root Ca and soil and root Mg. In contrast to these macronutrients, the micronutrients Fe and Zn and the toxic element Al decreased in the soil exchangeable fraction with the addition of wood ash, but not in the fine roots. Only Mn decreased in soil and in fine roots leading to a significant linear correlation between soil and root Mn. In soil, as well as in fine roots, strong positive correlations were found between the elements Ca and Mg and between Fe and Al. This indicates that the uptake of Mg resembles that of Ca and that of Al that of Fe. With the wood ash application, the pH increased from 3.2 to 4.8, the base saturation from 30% to 86%, the molar basic cations/Al ratio (BC/Al) of the soil solution from 1.5 to 5.5, and the molar Ca/Al ratio of the fine roots from 1.3 to 3.7. Overall, all below-ground indicators of soil acidification responded positively to the wood ash application within two years. Nitrate concentrations increased only slightly in the soil solution at a soil depth of 75–80 cm, and no signs of increased heavy metal concentrations in the soils or in the fine roots were apparent. This suggests that the recycling of wood ash could be an integral part of sustainable forest management because it closes the nutrient cycle and reverses soil acidification.     

Photosynthetic and Growth Response of Sugar Maple (Acer saccharum Marsh.) Mature Trees and Seedlings to Calcium,Magnesium, and Nitrogen Additions in the Catskill Mountains,NY, USA

Decline of sugar maple in North American forests has been attributed to changes in soil calcium (Ca) and nitrogen (N) by acidic precipitation. Although N is an essential and usually a limiting factor in forests, atmospheric N deposition may cause N-saturation leading to loss of soil Ca. Such changes can affect carbon gain and growth of sugar maple trees and seedlings. We applied a 2² factorial arrangement of N and dolomitic limestone containing Ca and Magnesium (Mg) to 12 forest plots in the Catskill Mountain region of NY, USA. To quantify the short-term effects, we measured photosynthetic-light responses of sugar maple mature trees and seedlings two or three times during two summers. We estimated maximum net photosynthesis (An-max) and its related light intensity (PAR at An-max), apparent quantum efficiency (A _qe), and light compensation point (LCP). To quantify the long-term effects, we measured basal area of living mature trees before and 4 and 8 years after treatment applications. Soil and foliar chemistry variables were also measured. Dolomitic limestone increased Ca, Mg, and pH in the soil Oe horizon. Mg was increased in the B horizon when comparing the plots receiving N with those receiving CaMg. In mature trees, foliar Ca and Mg concentrations were higher in the CaMg and N+CaMg plots than in the reference or N plots; foliar Ca concentration was higher in the N+CaMg plots compared with the CaMg plots, foliar Mg was higher in the CaMg plots than the N+CaMg plots; An-max was maximized due to N+CaMg treatment; A _qe decreased by N addition; and PAR at An-max increased by N or CaMg treatments alone, but the increase was maximized by their combination. No treatment effect was detected on basal areas of living mature trees four or eight years after treatment applications. In seedlings, An-max was increased by N+CaMg addition. The reference plots had an open herbaceous layer, but the plots receiving N had a dense monoculture of common woodfern in the forest floor, which can impede seedling survival.     

Controls of Bedrock Geochemistry on Soil and Plant Nutrients in Southeastern Utah

The cold deserts of the Colorado Plateau contain numerous geologically and geochemically distinct sedimentary bedrock types. In the area near Canyonlands National Park in Southeastern Utah, geochemical variation in geologic substrates is related to the depositional environment with higher concentrations of Fe, Al, P, K, and Mg in sediments deposited in alluvial or marine environments and lower concentrations in bedrock derived from eolian sand dunes. Availability of soil nutrients to vegetation is also controlled by the formation of secondary minerals, particularly for P and Ca availability, which, in some geologic settings, appears closely related to variation of CaCO₃ and Ca-phosphates in soils. However, the results of this study also indicate that P content is related to bedrock and soil Fe and Al content suggesting that the deposition history of the bedrock and the presence of P-bearing Fe and Al minerals, is important to contemporary P cycling in this region. The relation between bedrock type and exchangeable Mg and K is less clear-cut, despite large variation in bedrock concentrations of these elements. We examined soil nutrient concentrations and foliar nutrient concentration of grasses, shrubs, conifers, and forbs in four geochemically distinct field sites. All four of the functional plant groups had similar proportional responses to variation in soil nutrient availability despite large absolute differences in foliar nutrient concentrations and stoichiometry across species. Foliar P concentration (normalized to N) in particular showed relatively small variation across different geochemical settings despite large variation in soil P availability in these study sites. The limited foliar variation in bedrock-derived nutrients suggests that the dominant plant species in this dryland setting have a remarkably strong capacity to maintain foliar chemistry ratios despite large underlying differences in soil nutrient availability.