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.     

Assessment of the potential role of metals in sugar maple (Acer saccharum Marsh) decline in Ontario, Canada

Spatial and temporal differences in the crown condition of sugar maple (Acer saccharum Marsh) in Ontario remain largely unexplained. In this study, the potential role of metals in sugar maple dieback was explored by measuring metal concentrations in foliage and forest floor (LFH) at 35 forest stands in south-central Ontario that exhibit varying levels of decline and span a climatic, soil acidity and acid deposition gradient. Foliar and forest floor metal concentrations varied among sites by between two and ten-fold, with acidic sites exhibiting the highest concentrations of many metals in the forest floor and foliage. Sites with moderate decline symptoms (decline index (DI)?>?10, averaged between 1986 and 2004) had significantly greater Cd, Zn, Cu, Pb and Mn concentrations and lower Ca concentrations in the forest floor compared with healthy sites (DI?<?10). Foliar concentrations of Cd, Sr and Mn were also significantly greater and Ca was significantly lower in sites with moderate decline symptoms compared with healthy sites. However the highest metal concentrations in foliage and the forest floor found in this study are lower than critical values reported in the literature. The notable exception is Mn where values at acidic sites may be high enough to negatively impact sugar maple.     

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.     

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.     

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.     

Plant-soil feedback in spruce (Picea abies) and mixed spruce-beech (Fagus sylvatica) stands as indicated by dendrochemistry

Ten pairs of secondary pure spruce (Picea abies) and adjacent mixed spruce-beech (Fagus sylvatica) stands on comparable sites were selected on two different bedrocks for soil formation (Flysch: nutrient rich and high soil pH; Molasse: poor nutrient supply and acidic) to study how an admixture of beech to spruce stands affects nutrient cycling and consequently soil chemistry. Soil analyses indicated accumulation of Ca under the mixed stands while the top soil under pure spruce was acidified. It was hypothesized that changes of soil chemical properties due to species composition over the last six decades are reflected in the stem wood of spruce. Three healthy dominant spruce trees per plot were selected for coring. Cores were crossdated and half-decadal samples were analyzed for Ca, Mg, Mn and Al. Calcium and Mg concentrations in stem wood of spruce were significantly higher for the pure spruce than for the mixed stands in spite of lower Ca and Mg stores in the soil. We assume that acidification caused by pure spruce mobilized these cations temporarily, increasing soil solution contents and consequently stem wood concentrations. It was possible to reconstruct soil pH from the element ratios Ca/Al (pure stands) and Ca/Mg (mixed stands), since these ratios in the stem wood of the last half-decade did correlate with soil pH for selected soil depths. Reconstructed soil pH showed a decline over the last 60 years under both species compositions due to accumulation of base cations in the increasing biomass. Comparisons of reconstructed soil pH in 0–5 and 10–20 cm soil depth indicated more pronounced top soil acidification (lower soil pH in 0–5 cm) by spruce on the nutrient rich soil (Flysch) than on the acidic soil (Molasse). However, admixture of beech caused higher pH values in 0–5 cm than in 10–20 cm soil depth on Flysch due to the observed Ca-pump effect of beech (uptake of Ca from deeper soil horizons).     

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.     

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.     

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.     

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.     

The effect of regeneration burning upon the nutrient status of soil in two forest types in southern Tasmania

In two forest types in southern Tasmania, eucalypt rainforest (mixed forest) and eucalypt dry sclerophyll forest, surface soils (0–10 cm) from stands that had been clear-felled and burned between 1976 and 1979 were compared with those from uncut, unburned stands. Factors compared were total organic C, N, P, K, Mg, Ca, Zn, Mn; pH; exchangeable Ca, Mg, and K; cation exchange capacity; extractable P; soil phosphate buffering capacity; and N-mineralisation rates. Sampling started in April 1979 and ended in October 1980. Within each forest type, soils from burned coupes had higher mean values for pH, exchangeable cations, percent base saturation, and nitrate-N produced during aerobic incubation, and had lower mean values for exchangeable acidity and ammonium-N produced during aerobic incubation than soils from unburned coupes. In mixed forest only, soils from burned coupes had higher mean values for extractable P and soil phosphate buffering capacity, and lower mean values for total organic C than those of unburned coupes. There were only small differences between burned and unburned soils in cation exchange capacity and ammonium-N produced during anaerobic incubation. For each burned coupe in mixed forest, with increase in time since burning there was a decrease in pH, an increase in exchangeable acidity, and a decrease in rate of production of nitrate: no changes were detected in other factors. It is concluded that, for clay soils developed on dolerite, the nutritional status of soil in both forest types is probably improved by burning. The improvement lasts for more than 4 years in mixed forest and more than two years in dry sclerophyll forest. Only minor leaching of nutrients to below 10 cm in depth is likely to occur in either type.     

Does soil acidity reduce subsoil rooting in Norway spruce (Picea abies)?

Increasing evidence suggests that forest soils in central and northern Europe as well as in North America have been significantly acidified by acid deposition during the last decades. The present investigation was undertaken to examine the effect of soil acidity on rooting patterns of 40-year-old Norway spruce trees by comparing fine and coarse roots among four stands which differed in soil acidity and Mg (and Ca) nutrition. The coarse root systems of four to five 40-year-old Norway spruce trees per stand were manually excavated. The sum of cross sectional area (CSA) at 60 cm soil depth and below of all vertical coarse roots, as a measure of vertical rooting intensity, was strongly reduced with increasing subsoil acidity of the stands. This pattern was confirmed when 5 additional acidic sites were included in the analysis. Fine root biomass in the mineral soil estimated by repeated soil coring was strongly reduced in the heavily acidified stands, but increased in the humic layer. Using ingrowth cores and a screen technique, we showed that the higher root biomass in the humic layer of the more acidic stands was a result of higher root production. Thus, reduced fine root biomass and coarse root CSA in deeper soil layers coincided with increased root growth in the humic layer. Root mineral analysis showed Ca/Al ratios decreased with decreasing base saturation in the deeper mineral soil (20–40 cm). In the top mineral soil, only minor differences were observed among stands. In general, low Ca/Al ratios coincided with low fine root biomass. Calcium/aluminum ratios determined in cortical cell walls using X-ray microanalysis showed a similar pattern as Ca/Al ratios based on analysis of whole fine roots, although the amplitude of changes among the stands was much greater. Aluminum concentrations and Ca/Al ratios in cortical cell walls were at levels found to inhibit root growth of spruce seedlings in laboratory experiments. The data support the idea that Al (or Ca/Al ratios) and acid deposition-induced Mg (and possibly Ca) deficiency are important factors influencing root growth and distribution in acidic forest soils. Changes in carbon partitioning within the root system may contribute to a reduction in deep root growth.     

Wood ash applications to temperate forest ecosystems—potential benefits and drawbacks

The objectives of the present work were (a) to quantify the effects of wood ash on forest ecosystems through a meta-analysis approach associated with a detailed review of the literature (mainly composed of work carried out in Nordic countries) and (b) to extrapolate the effects on forest growth to other contexts (i.e. warm temperate countries) by identifying the cases for which wood ash applications can be beneficial to forest production. Three databases were built regarding the effects of wood ash on soil (151 observations; 33 experimental field trials), on nutrient concentrations of tree foliage (68 observations; 28 trials) and on annual stem biomass growth rate (70 observations; 27 trials). We obtained information on the wide variability of ash properties due to differences in burnt compounds, combustion processes and ash conditioning. Two important properties of wood ash are its high pH value and neutralizing capacity. These properties result in biochemical modifications of forest soils limed with ash. In the short term, soil solution composition was dramatically modified. Intense peaks of the K, Na or SO₄ concentrations were observed, resulting from the dissolution of salts contained in ash. At the same time, Ca and Mg concentrations increased as the carbonate pool of wood ash started to dissolve. The consequence of this dissolution process was an increase in the pH in all the soil phases. These modifications increased the activity of the soil microflora and some isolated peaks in the mineralization of soil organic matter may be observed in mineral soils. In the longer term, that is to say after the first year following ash application, only the effect on the acidity status of the soil remained significant. The effects of ash addition on forest ecosystems usually increased with the dose and were more pronounced with loose ash compared to aggregated ash. The addition of wood ash into forest ecosystems increased the foliar Ca status of trees. Some modifications of other nutrients, like P or K, were also observed but only for a few years after treatment. For most stands growing on mineral soils of Nordic countries, this treatment did not result in an increase in tree growth, probably because of the absence of N in the ash. For stands growing on organic soils of the same area, this input, associated with a long-lasting increase of soil organic matter mineralization, was sufficient to improve tree growth significantly (median = +59% compared to the control). For soils located in warm temperate regions, similar responses are expected for organic soils. For mineral soils, the wood ash application is expected to be suitable for stands showing deficiencies in K, Ca or Mg. Ash may contain high amounts of toxic heavy metals such as Cd. The bioavailability of most of these elements appeared to be very low in a forest context. No contamination of food chains has been observed, except possibly via some species of fungi, and heavy metals remain in the forest litter or in the topsoil. Based on all the reviewed results, several guidelines for wood ash application into forest ecosystems are proposed. Wood ash application should be restricted to acidic soils. Applications should consist of low doses of a stabilized ash form. Wood ash should be applied to adult stands rather than onto seedlings.     

Aluminium and manganese and their relation to calcium in soil solution and needles in three Norway spruce (Picea abies,L. Karst.) stands of Upper Austria

Soil solution and needles of three mature spruce stands in Upper Austria were analysed in order to investigate the uptake and possible toxic effects of Mn and Al, as these two elements become highly mobilised in the soil due to increasing acidity. The Ca/Al molar ratio in the soil solution was below 0.2 in the most damaged stand during almost the whole vegetation period. Despite different dynamics, Al reaches almost identical values in all stands at the end of the vegetation period in both 1-year (current) and 2-year-old needles, respectively. Therefore, needle analysis is not a useful tool for estimation of free Al in the soil. Needle contents of other elements could provide a better information for understanding the forest decline. Mn in the needles correlates significantly with Mn concentrations in the soil solution. As soil Mn will be mobilised by acidic input, Mn needle content can increase to very high levels. Manganese distribution, its interaction with calcium, and possible toxic effects are discussed.     

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.     

Nutrient cycling and foliar status in an urban pine forest in Athens, Greece

The nutrient cycling and foliar status for the elements Ca, Mg, K, N, P, S, Fe, Mn, Zn and Cu were investigated in an urban forest of Aleppo pine (Pinus halepensis) in 2004 in Athens, Greece in order to draw conclusions on the productivity status and health of the ecosystem. The fluxes of bulk and throughfall deposition were characterized by the high amounts of Ca, organic N and sulfate S. The magnitude of the sulfate S fluxes indicated a polluted atmosphere. The nutrient enrichment in throughfall was appreciable for ammonium N, P and Mn. The mineral soil formed the largest pool for all the elements followed by the forest floor, trunk wood and trunk bark. The understory vegetation consisting of annual plants proved important for storing N, P and K. Compared to current year needles of Aleppo pine in remote forests of Spain, the needles of the Aleppo pine trees in Athens had significantly higher concentrations of Ca, N, P and Cu and significantly lower concentrations of Mg and Zn. The soil had a high concentration of calcium carbonate and accordingly high pH values. When all inputs to the forest floor were taken into account, the mean residence time of nutrients in the forest floor followed the order Fe > Mn > Cu > Ca > Mg > P > Zn > N > K > S.     

Effects of wood ash and nitrogen fertilization on fine root biomass and soil and foliage nutrients in a Norway spruce stand in Finland

We determined the effects of wood ash fertilization, given together with nitrogen (WAN), and nitrogen given together with P, B and Cu (SSF), on soil and foliage nutrients and fine root biomass in a 45-year-old Norway spruce stand in southern Finland. Fine roots were sampled 9 years, and the soil 10 years after ash (3 t/ha) and nitrogen (150 kg/ha) application. Fine root biomass tended to be lower, the necromass higher, and the fine root distribution relatively deeper on the WAN than on the control and SSF plots. The response of fine root biomass to WAN was probably related to changes in soil acidity. pH, base saturation, total and extractable concentrations of Ca, K, Mg and P, and total B, Cd, Mn, Ni and Zn concentrations in the organic layer were significantly higher on the WAN plots than on the SSF and the control plots with no ash and nutrient addition. On the WAN plots, the pH was 1.2 pH-units higher, the exchangeable Ca concentrations fourfold and those of Mg over twofold compared to the control plots. WAN increased the concentrations of K but decreased those of Mn and Ni in the needles compared to the control and SSF treatment. Even though ash and nitrogen fertilisation tended to decrease the fine root biomass, this decrease was not likely to affect tree growth during a 10-year period.     

Nutrient transfer by leaf litterfall during a sugar maple decline episode at Lake Clair watershed,Québec,Canada

In a declining sugar maple (SM) stand, we tested the hypothesis that an increasing relative abundance of American beech (AB) and yellow birch (YB) would improve litter quality by providing a higher proportion of litterfall richer in base cations and lower in acidity. From 1989 to 2006, SM leaf fall diminished from 59% (1,718 kg ha⁻¹ year⁻¹) to 36% (915 kg ha⁻¹ year⁻¹) of the total leaf fall biomass. Overall, the increase in AB and YB litterfall compensated for the SM decrease, resulting in constant annual leaf litterfall fluxes (2,803 kg ha⁻¹ year⁻¹) over the period studied. However, because the leaf litter for AB and YB had Ca and Mg concentrations 2–3 times higher than did SM, Ca and Mg concentrations and fluxes in leaf litterfall significantly increased between 1989 and 2006. Leaf litterfall of AB and YB also has a higher base/acid ratio than SM. Consequently, changes in forest composition following SM decline led to a clear improvement in litterfall quality in terms of base cations content and fluxes and acid–base properties.     

不同季节亚热带常绿阔叶林6个树种凋落叶钙、镁、锰的释放特征

采用凋落物分解袋法,研究了亚热带常绿阔叶林区马尾松、柳杉、杉木、香樟、红椿、麻栎6个典型树种凋落叶的Ca、Mg、Mn在第一个分解年不同雨热季节的释放特征.结果表明：经历1年的分解,6种凋落叶Ca、Mg、Mn释放率分别为-13.8%～92.3%、4.0%～64.8%和41.6%～81.1%.马尾松和香樟凋落叶Ca释放动态呈现富集-释放模式,其余4种凋落叶整体上呈现释放的模式;香樟凋落叶Mg释放动态呈现富集-释放模式,其余5种凋落叶呈现直接释放模式;柳杉和红椿凋落叶Mn释放动态呈现富集-释放模式,其余4种凋落叶呈现直接释放模式.凋落叶Ca、Mg、Mn的释放明显受到季节性降雨的影响,且因凋落叶种类不同而有差异.Ca、Mg、Mn在雨季的释放率和释放量均高于旱季.初始养分含量和降雨量是影响凋落物分解过程元素释放的重要因子.     

Fine Root Dynamics and Forest Production Across a Calcium Gradient in Northern Hardwood and Conifer Ecosystems

Losses of soil base cations due to acid rain have been implicated in declines of red spruce and sugar maple in the northeastern USA. We studied fine root and aboveground biomass and production in five northern hardwood and three conifer stands differing in soil Ca status at Sleepers River, VT; Hubbard Brook, NH; and Cone Pond, NH. Neither aboveground biomass and production nor belowground biomass were related to soil Ca or Ca:Al ratios across this gradient. Hardwood stands had 37% higher aboveground biomass (P = 0.03) and 44% higher leaf litter production (P < 0.01) than the conifer stands, on average. Fine root biomass (<2 mm in diameter) in the upper 35 cm of the soil, including the forest floor, was very similar in hardwoods and conifers (5.92 and 5.93 Mg ha⁻¹). The turnover coefficient (TC) of fine roots smaller than 1 mm ranged from 0.62 to 1.86 y⁻¹ and increased significantly with soil exchangeable Ca (P = 0.03). As a result, calculated fine root production was clearly higher in sites with higher soil Ca (P = 0.02). Fine root production (biomass times turnover) ranged from 1.2 to 3.7 Mg ha⁻¹ y⁻¹ for hardwood stands and from 0.9 to 2.3 Mg ha⁻¹ y⁻¹ for conifer stands. The relationship we observed between soil Ca availability and root production suggests that cation depletion might lead to reduced carbon allocation to roots in these ecosystems.