Evaluating, and possibly ameliorating, the effects of base cation depletion in forest soils caused by acid deposition is an important topic in the northeastern United States. We added 850 kg Ca ha-¹ ...as wollastonite (CaSiO₃) to an 11.8-ha watershed at the Hubbard Brook Experimental Forest (HBEF), a northern hardwood forest in New Hampshire, USA, in fall 1999 to replace calcium (Ca) leached from the ecosystem by acid deposition over the past 6 decades. Soil microbial biomass carbon (C) and nitrogen (N) concentrations, gross and potential net N mineralization and nitrification rates, soil solution and stream chemistry, soil:atmosphere trace gas (CO₂, N₂O, CH₄) fluxes, and foliar N concentrations have been monitored in the treated watershed and in reference areas at the HBEF before and since the Ca addition. We expected that rates of microbial C and N cycle processes would increase in response to the treatment. By 2000, soil pH was increased by a full unit in the Oie soil horizon, and by 2002 it was increased by nearly 0.5 units in the Oa soil horizon. However, there were declines in the N content of the microbial biomass, potential net and gross N mineralization rates, and soil inorganic N pools in the Oie horizon of the treated watershed. Stream, soil solution, and foliar concentrations of N showed no response to treatment. The lack of stimulation of N cycling by Ca addition suggests that microbes may not be stimulated by increased pH and Ca levels in the naturally acidic soils at the HBEF, or that other factors (for example, phosphorus, or Ca binding of labile organic matter) may constrain the capacity of microbes to respond to increased pH in the treated watershed. Possible fates for the approximately 10 kg N ha-¹ decline in microbial and soil inorganic pools include components of the plant community that we did not measure (for example, seedlings, understory shrubs), increased fluxes of N₂ and/or N storage in soil organic matter. These results raise questions about the factors regulating microbial biomass and activity in northern hardwood forests that should be considered in the context of proposals to mitigate the depletion of nutrient cations in soil.
The effects of acidic deposition in the northeastern US include the acidification of soil and water, which stresses terrestrial and aquatic biota. Driscoll et al examine the ecological effects of ...acidic deposition in New England and New York and explore the relationship between emissions reductions and ecosystem recovery.
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BFBNIB, DOBA, IZUM, KILJ, NMLJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
The depletion of calcium in forest ecosystems of the northeastern USA is thought to be a consequence of acidic deposition and to be at present restricting the recovery of forest and aquatic systems ...now that acidic deposition itself is declining. This depletion of calcium has been inferred from studies showing that sources of calcium in forest ecosystems-namely, atmospheric deposition and mineral weathering of silicate rocks such as plagioclase, a calcium-sodium silicate-do not match calcium outputs observed in forest streams. It is therefore thought that calcium is being lost from exchangeable and organically bound calcium in forest soils. Here we investigate the sources of calcium in the Hubbard Brook experimental forest, through analysis of calcium and strontium abundances and strontium isotope ratios within various soil, vegetation and hydrological pools. We show that the dissolution of apatite (calcium phosphate) represents a source of calcium that is comparable in size to known inputs from atmospheric sources and silicate weathering. Moreover, apatite-derived calcium was utilized largely by ectomycorrhizal tree species, suggesting that mycorrhizae may weather apatite and absorb the released ions directly, without the ions entering the exchangeable soil pool. Therefore, it seems that apatite weathering can compensate for some of the calcium lost from base-poor ecosystems, and should be considered when estimating soil acidification impacts and calcium cycling.
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DOBA, IJS, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
Red spruce (Picea rubens Sarg.) trees are uniquely vulnerable to foliar freezing injury during the cold season (fall and winter), but are also capable of photosynthetic activity if temperatures ...moderate. To evaluate the influence of calcium (Ca) addition on the physiology of red spruce during the cold season, we measured concentrations of foliar polyamines and free amino acids (putative stress-protection compounds), chlorophyll (a key photosystem component), and sapwood area (a proxy for foliar biomass), for trees in Ca-addition (CaSiO3 added) and Ca-depleted (reference) watersheds at the Hubbard Brook Experimental Forest (NH, USA). Ca-addition increased concentrations of the amino acids alanine and γ-aminobutyric acid (GABA) and the polyamines putrescine (Put) and spermidine (Spd) in November, and Put in February relative to foliage from the reference watershed. Consistent with increased stress protection, foliage from the Ca-addition watershed had higher total chlorophyll and chlorophyll a concentrations in February than foliage from the reference watershed. In contrast, foliage from the reference watershed had significantly lower glutamic acid (Glu) and higher alanine (Ala) concentrations in February than foliage from the Ca-addition watershed. Imbalances in Ala:Glu have been attributed to cold sensitivity or damage in other species. In addition to concentration-based differences in foliar compounds, trees from the Ca-addition watershed had higher estimated levels of foliar biomass than trees from the reference watershed. Our findings suggest that Ca-addition increased the stress tolerance and productive capacity of red spruce foliage during the cold season, and resulted in greater crown mass compared to trees growing on untreated soils.
To assess potential forest compositional responses to exchangeable soil calcium (Ca(exch)) and aluminum (Al(exch)), we characterized light-dependent growth and mortality of tree seedlings under ...amendments of CaCl2 and AlCl3 at Hubbard Brook Experimental Forest (HBEF), New Hampshire, U.S.A. Seedlings of Acer saccharum Marsh., Fagus grandifolia Ehrh., Betula alleghaniensis Britton, Abies balsamea (L.) Mill., and Picea rubens Sarg. were transplanted into field plots, which were randomly assigned to control, CaCl2, or AlCl3 treatments and stratified across <1 to 35% full sun. Acer saccharum and P. rubens exhibited significantly higher mortality in Al-amended than Ca-amended or control plots. Acer saccharum showed significant increases in relative diameter growth in Ca-amended plots versus controls; all other species showed nonsignificantly higher relative diameter growth under Ca amendments. We incorporated significant seedling responses into a model of forest dynamics (SORTIE) to assess potential changes in species composition under Al(exch) increases and Ca(exch) losses. SORTIE predicts that further increases in Al(exch) would have negligible effects on canopy composition within 200 years but that the estimated Ca(exch) depleted from HBEF between 1968 and 1995 and its influence on seedling dynamics could lead to substantial decreases in A. saccharum canopy dominance within a single forest generation (<125 years).
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BF, DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
Watershed budget studies at the Hubbard Brook Experimental Forest (HBEF), New Hampshire, USA, have demonstrated high calcium depletion of soil during the 20th century due, in part, to acid ...deposition. Over the past 25 years, tree growth (especially for sugar maple) has declined on the experimental watersheds at the HBEF. In October 1999, 0.85 Mg Ca/ha was added to Watershed 1 (W1) at the HBEF in the form of wollastonite (CaSiO3), a treatment that, by summer 2002, had raised the pH in the Oie horizon from 3.8 to 5.0 and, in the Oa horizon, from 3.9 to 4.2. We measured the response of sugar maple to the calcium fertilization treatment on W1. Foliar calcium concentration of canopy sugar maples in W1 increased markedly beginning the second year after treatment, and foliar manganese declined in years four and five. By 2005, the crown condition of sugar maple was much healthier in the treated watershed as compared with the untreated reference watershed (W6). Following high seed production in 2000 and 2002, the density of sugar maple seedlings increased significantly on W1 in comparison with W6 in 2001 and 2003. Survivorship of the 2003 cohort through July 2005 was much higher on W1 (36.6%) than W6 (10.2%). In 2003, sugar maple germinants on W1 were approximately 50% larger than those in reference plots, and foliar chlorophyll concentrations were significantly greater (0.27 g/m2 vs. 0.23 g/m2 leaf area). Foliage and fine-root calcium concentrations were roughly twice as high, and manganese concentrations twice as low in the treated than the reference seedlings in 2003 and 2004. Mycorrhizal colonization of seedlings was also much greater in the treated (22.4% of root length) than the reference sites (4.4%). A similar, though less dramatic, difference was observed for mycorrhizal colonization of mature sugar maples (56% vs. 35%). These results reinforce and extend other regional observations that sugar maple decline in the northeastern United States and southern Canada is caused in part by anthropogenic effects on soil calcium status, but the causal interactions among inorganic nutrition, physiological stress, mycorrhizal colonization, and seedling growth and health remain to be established.
▶ Ca addition to forest soil caused species-specific changes in foliar metabolism. ▶ Of the three species, maple was the most responsive followed by birch. ▶ Higher elevation trees responded better ...than lower elevation trees. ▶ Indicators of physiological stress, i.e., putrescine, arginine and GABA decreased. ▶ Foliar soluble Ca, P, chlorophyll, glutamate and glycine increased.
Acidic deposition has caused a depletion of calcium (Ca) in the northeastern forest soils. Wollastonite (Ca silicate) was added to watershed 1 (WS1) at the Hubbard Brook Experimental Forest (HBEF) in 1999 to evaluate its effects on various functions of the HBEF ecosystem. The effects of Ca addition on foliar soluble (extractable in 5% HClO
4) ions, chlorophyll, polyamines, and amino acids were studied in three hardwood species, namely sugar maple, yellow birch, and American beech. We further analyzed these effects in relation to elevation at Ca-supplemented WS1 and reference WS3 watersheds. Foliar soluble Ca increased significantly in all species at mid and high elevations at Ca-supplemented WS1. This was accompanied by increases in soluble P, chlorophyll, and two amino acids, glutamate and glycine. A decrease in known metabolic indicators of physiological stress (i.e., the amino acids, arginine and γ-aminobutyric acid (GABA), and the diamine, putrescine) was also observed. In general, these changes were species-specific and occurred in an elevation dependent manner. Despite an observed increase in Ca at high elevation for all three species, only sugar maple exhibited a decrease in foliar putrescine at this elevation indicating possible remediation from Ca deficiency. At higher elevations of the reference WS3 site, foliar concentrations of Ca and Mg, as well as Ca:Mn ratios were lower, whereas Al, putrescine, spermidine, and GABA were generally higher. Comparison of metabolic data from these three species reinforces the earlier findings that sugar maple is the most sensitive and American beech the least sensitive species to soil Ca limitation. Furthermore, there was an increase in sensitivity with an increase in elevation.
We surveyed and wounded forest-grown sugar maple (Acer saccharum Marsh.) trees in a long-term, replicated Ca manipulation study at the Hubbard Brook Experimental Forest in New Hampshire, USA. Plots ...received applications of Ca (to boost Ca availability above depleted ambient levels) or Al (to compete with Ca uptake and further reduce Ca availability). We found significantly greater total foliar and membrane-associated Ca in foliage of trees in plots fertilized with Ca when compared with trees from Al-addition and control plots (P = 0.005). Coinciding with foliar Ca differences, trees exhibited a significant difference in crown vigor and in percent branch dieback among treatments (P < 0.05), with a trend towards improved canopy health as Ca levels increased. Annual basal area increment growth for the years following treatment initiation (1998-2004) was significantly greater in trees subjected to Ca addition compared with trees in control and Al treatments. Treatment-related improvements in growth were particularly evident after overstory release following a 1998 ice storm. The amount of wound closure was also greatest for trees in Ca-addition plots relative to Al-addition and control plots (P = 0.041). These findings support evidence that ambient Ca depletion is an important limiting factor regarding sugar maple health and highlight the influence of Ca on wound closure and growth following release from competition.
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Dostopno za:
BF, DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
In fall (November 2005) and winter (February 2006), we collected current-year foliage of native red spruce (Picea rubens Sarg.) growing in a reference watershed and in a watershed treated in 1999 ...with wollastonite (CaSiO3, a slow-release calcium source) to simulate preindustrial soil calcium concentrations (Ca-addition watershed) at the Hubbard Brook Experimental Forest (Thornton, NH). We analyzed nutrition, soluble sugar concentrations, ascorbate peroxidase (APX) activity and cold tolerance, to evaluate the basis of recent (2003) differences between watersheds in red spruce foliar winter injury. Foliar Ca and total sugar concentrations were significantly higher in trees in the Ca-addition watershed than in trees in the reference watershed during both fall (P = 0.037 and 0.035, respectively) and winter (P = 0.055 and 0.036, respectively). The Ca-addition treatment significantly increased foliar fructose and glucose concentrations in November (P = 0.013 and 0.007, respectively) and foliar sucrose concentrations in winter (P = 0.040). Foliar APX activity was similar in trees in both watersheds during fall (P = 0.28), but higher in trees in the Ca-addition watershed during winter (P = 0.063). Cold tolerance of foliage was significantly greater in trees in the Ca-addition watershed than in trees in the reference watershed (P < 0.001). Our results suggest that low foliar sugar concentrations and APX activity, and reduced cold tolerance in trees in the reference watershed contributed to their high vulnerability to winter injury in 2003. Because the reference watershed reflects forest conditions in the region, the consequences of impaired physiological function caused by soil Ca depletion may have widespread implications for forest health.
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