15N标记氮在盆栽山毛榉中的分配

温室条件下,向盆栽山毛榉幼苗中施加192g.m·^-2^15N示踪物,研究连续两个生长季沉积氮在森林土壤（含森林地被物）、沥出物、以及地上和地下部分生物量的分配。模拟了四种处理（栽培和非栽培）下的NH4^＋和NO3^-沉积情况,每种处理各自标记为^15N-NH4^＋或者^15N-NO3^-。在整个体系中施加15N的总回收率分别是,盆栽处理的^15N-NH4^＋为67.3%～74.9%,非盆栽处理的^15N-NO3^-为85.3%～88.1%。两种^15N示踪物主要沉积在森林土壤（包括森林地被物）中,其中盆栽处理的森里土壤中^15N-NH4^＋为34.6%～33.7%,^15N-NO3^-为13.1%～9.0%,说明异养微生物有很强的固氮作用。森林土壤微生物对^15N-NH4^＋的固定能力比^15N-NO3^-的固定能力强三倍。^15N-NH4^＋的优先异养利用造成土壤中^15N-NH4^＋的沉积量是植物体保存量的两倍 而土壤中^15N-NO3^-的沉积量却低于植物体的。总之,植被-土壤系统中15N-NH4＋的沉积量比^15N-NO3^-的沉积高了60%,说明了沉积氮的形式在森林生态系统中氮保存中的重要性。     

Litter impair spruce seedling emergence in beech forests: a litter manipulation experiment

European beech Fagus sylvatica and Norway spruce Picea abies are economically and ecologically important forest trees in large parts of Europe. Today, the beech forest reaches its northern distribution limit in south-eastern Norway and it is expected to expand northwards due to climate warming. This expansion will likely result in fundamental ecosystem changes. To increase our knowledge about the competitive balance between spruce and beech, we have investigated how beech and spruce litter affect spruce seedling emergence, growth and uptake of C and N. We did this in a seed-sowing experiment that included litter layer removal as well as reciprocal transplantations of litter layers between spruce and beech forests. Our results show that spruce seedling emergence was significantly impaired by both litter layer types, and especially so by the beech litter layer in the beech forest. The low seedling emergence in beech forests is concurrent with their lower light availability.     

Identification of viruses from European beech (Fagus sylvatica L.) of declining forests in Northrhine-Westfalia (FRG)

Five different viruses isolated from degenerating European beech of different forest sites of West Germany could be identified as cherry leaf roll virus, brome mosaic virus, potex and potyviruses.     

Leaf and litter nitrogen and phosphorus in three forests with low P supply

We compared the N and P contents of the main labile components of nutrient cycles in three different forest ecosystems [a tropical evergreen forest (TEF); a tropical dry forest (TDF); and a Mediterranean temperate forest (MTF)] with low P supply. A mass-balance approach was used to estimate mean residence times for organic matter, N and P in the forest floor, and to examine the flexibility of N and P intra-system cycling in the three forest ecosystems. For this purpose, we combined published values of N and P in foliage, litterfall, forest floor litter and mineral soils in these three forest ecosystems. The results of our analysis were consistent with the widely held belief that the N content of leaves (both green and senescent) and litter increases with increasing temperatures. In contrast, the data did not support the hypothesis that leaf P content decreases with increasing temperatures and precipitation: leaf and litterfall P contents were higher in both tropical forests than they were in the temperate forest. The TEF had the highest P content of the three forests studied. The mass-balance analysis indicated that although P mineralization in the TDF can run ahead of litter decomposition stoichiometry when P is in short supply, flexibility is much reduced or absent in the TEF and the MTF. Our analysis provides additional evidence of the importance of climatic factors in forest ecosystem processes and highlights the role of flexibility in ecosystem nutrient cycling, especially for P in ecosystems with a limited P supply.     

Effects of litter quality and parent material on organic matter characteristics and N-dynamics in Luxembourg beech and hornbeam forests

To test effects of litter quality and soil conditions on N-dynamics, we selected seven forests in Luxembourg dominated by beech (Fagus sylvatica, L.) and hornbeam (Carpinus betulus L.), and located on acid loam, decalcified marl or limestone, and measured organic matter characteristics, microbial C and N and net N-mineralization in a laboratory incubation experiment. Organic layer characteristics were significantly affected by species, with lower litter decay and higher accumulation under the less palatable beech, even on limestone. However, beech and hornbeam did not show any differences in N-cycling at all. Instead of species, N-cycling was affected by site conditions, albeit different than expected. Microbial N generally increased from acid loam to limestone, but acid loam showed higher net N-mineralization, especially in the organic layer. Also, acid loam showed high instead of low efficiency of N-mineralization per unit microbe, in both organic layer and mineral topsoil. In addition, acid loam showed net consumption of DOC instead of release in both soil layers, which suggests that not N, but C was a limiting factor to decomposition. In contrast, limestone showed low net N-mineralization in the organic layer, despite high mass and well-decomposed organic matter, and low efficiency of N-mineralization per unit microbe in both organic layer and mineral topsoil. DOC was net released instead of consumed, which supported that not C, but N was a limiting factor. The general lack of differences in net N-cycling between species, but relatively clear site effects, is discussed in relation to different microbial strategies. Acid soil may have high net N-release despite low biological activity, because N-requirements of fungi are also low, while in calcareous soil, high bacterial N-demand may counteract high gross N-release. Thus, species producing litter that decomposes rapidly may be planted to improve soil conditions and plant biodiversity, but litter quality effects on N-availability may be less important than soil conditions.     

Climate and forest management influence nitrogen balance of European beech forests: microbial N transformations and inorganic N net uptake capacity of mycorrhizal roots

The effects of local climate and silvicultural treatment on the inorganic N availability, net N uptake capacity of mycorrhizal beech roots and microbial N conversion were assessed in order to characterise changes in the partitioning of inorganic N between adult beech and soil microorganisms. Fine root dynamics, inorganic N in the soil solution and in soil extracts, nitrate and ammonium uptake kinetics of beech as well as gross ammonification, nitrification and denitrification rates were determined in a beech stand consisting of paired sites that mainly differed in aspect (SW vs. NE) and stand density (controls and thinning treatments). Nitrate was the only inorganic N form detectable in the soil water. Its concentration was high in control plots of the NE aspect, but only in canopy gaps and not influenced by thinning. Neither thinning nor aspect affected the abundance of root tips in the soil. Maximum nitrate net uptake by mycorrhizal fine roots of beech, however, differed with aspect, showing significantly lower values at the SW aspect with warm–dry local climate. There were no clear-cut significant effects of local climate or thinning on microbial N conversion, but a tendency towards higher ammonification and nitrification and lower denitrification rates on the untreated controls of the SW as compared to the NE aspect. Apparently, the observed sensitivity of beech towards reduced soil water availability is at least partially due to impaired N acquisition. This seems to be mainly a consequence of reduced N uptake capacity rather than of limited microbial re-supply of inorganic N or of changed patterns of inorganic N partitioning between soil bacteria and roots.     

Habitat factors for land snails in European beech forests with a special focus on coarse woody debris

Analyses of land snails and habitat factors in acid beech forests were conducted in southern Germany (northern Bavaria). The objectives were to study the effects of habitat characteristics on snail density and species richness. Habitat structures were determined for 37 plots in one big forest. We found a significant relationship between the number of snail species and individuals and the following set of habitat factors coverage of herbaceous layer, growing stock, mean diameter at breast height of the three largest trees (DBHmax), stand age, total dead wood volume per ha, and advanced decomposed dead wood volume per ha. We use maximally selected rank statistics to estimate cutpoints separating stands with low densities, from stands with high snail densities. Here, we define cutpoints for a significant higher snail density at a stand age of 187 years, 57 m³/ha dead wood, 40 m³/ha advanced decomposed dead wood, 63 cm DBHmax and more than 1% herbaceous layer. For species richness, cutpoints are estimated at 338 m³/ha stand volume, 170 years stand age, 50 m³/ha total dead wood amount, 15 m³/ha advanced decomposed dead wood and 56 cm DBHmax. The microhabitat analysis shows a higher pH value and a higher Calcium content at the bottom of large snags and under large lying dead wood pieces in comparison to litter, upper mineral soil and at the bottom of vital living trees. Snail species and individual density are significantly linked to these patterns of chemical parameters. The identified cutpoints are a good base for ecological management decisions in forest management.     

Characterization of Cylindrocarpon‐like species associated with litter in the old‐growth beech forests of Central Europe

Cylindrocarpon‐like fungi are globally distributed plant pathogens and have a wide range of host species. However, very little is known about the species that live in the topsoil of forests and their potential role in reducing the natural regeneration of tree species, particularly of forest trees that produce abundant fruit only once every few years. To enhance our understanding of the species that inhabit the topsoil, we studied the diversity and pathogenicity of Cylindrocarpon‐like fungi inhabiting the litter in old‐growth mixed‐beech forests in the Carpathians (Poland) and in the Alps (Austria), and in a managed beech stand in the Krakowsko‐Cz?stochowska Highland (Poland). The fungi inhabiting the beech litter were investigated using beechnuts and pine seedlings as bait. Isolates were identified based on morphology and DNA sequencing. The pathogenicity of the most common species was investigated by inoculating beech germinants. A wide range of Cylindrocarpon‐like fungi were associated with the beech litter: 718 cultures representing 12 species were isolated. Five taxa were identified down to species level: namely Ilyonectria crassa, I. pseudodestructans, I. rufa, Neonectria candida and N. obtusispora, and seven species were identified to genus level (Neonectria or Ilyonectria species). Ilyonectria destructans, which is considered to be the sexual morph of ‘Cylindrocarpon destructans’, was not found. There were qualitative and quantitative differences between the different forest sites in terms of Neonectria and Ilyonectria species composition in beech litter. The isolation frequency and species richness of Cylindrocarpon‐like fungi were greatest in beech litter taken from old‐growth mixed‐beech forests. Neonectria and Ilyonectria species were capable of killing beech germinants, suggesting that they may play a negative role in natural beech regeneration.     

Decomposition of needle/leaf litter from Scots pine, black cherry, common oak and European beech at a conurbation forest site

Litter decomposition was studied for 2 years in a mixed forest serving as a water protection area (Rhine-Neckar conurbation, SW Germany). Two experiments differing in initial dry weight equivalent in litterbags were set up: one to compare decomposition of European beech leaves (Fagus sylvatica) with common oak leaves (Quercus robur), and the other comparing decomposition of Scots pine needles (Pinus sylvestris) with black cherry leaves (Prunus serotina Ehrh.), respectively. Mass losses were greater for oak litter than for beech (75.0 versus 34.6%), and for cherry litter than for pine (94.6 versus 68.3%). In both experiments, a strong initial loss of soluble compounds occurred. The changes in litter N and P concentrations and the decrease in C-to-N ratio coincided with changes in residual mass. However, neither tannin and phenolic concentrations nor NMR could explain the pronounced variation in mass loss after 2 years. Differences in litter palatability and toughness, nutrient contents and other organic compounds may be responsible for the considerable differences in residual mass between litter types. The fast decay of black cherry leaves appears to play a major role in the present humus dynamics at the studied site. Since black cherry has a high N demand, which is mainly met by root uptake from the forest floor, this species is crucial for internal N cycling at this conurbation forest site. These effects together may significantly contribute to prevent nitrate leaching from the forest ecosystem which is subject to a continuous N deposition on an elevated level.     

Competition for nitrogen between adult European beech and its offspring is reduced by avoidance strategy

Plant growth, reproduction, and biomass allocation may be affected differently by nitrogen availability depending on tree size and age. In this context, competition for limited N may be avoided by different strategies of N acquisition between different vegetation components (i.e., seedlings, mature trees, other woody and herbaceous understorey). This study investigated in a field experiment whether the competition for N between different vegetation components in beech forests was prevented via seasonal timing of N uptake and affected by microbial N use. For this purpose, a removal approach was used to study the seasonal effects on N uptake and N metabolites in adult beech trees and beech natural regeneration, as well as soil microbial processes of inorganic N production and utilisation. We found that the competition for N between beech natural regeneration and mature beech trees was reduced by seasonal avoidance strategies (“good parenting”) of N uptake regardless of the N sources used. In spring, organic and inorganic N uptake capacity was significantly higher in beech seedlings compared to adult beech trees, whereas in autumn mature beech trees showed the highest N uptake rates. Removal of vegetation components did not result in changes in soil microbial N processes in the course of the growing season. Thus, N resources released by the removal of vegetation components were marginal. This consistency in soil microbial N processes indicates that competition between plants and soil microorganisms for N was not avoided by timing of acquisition during the vegetation period, but existed during the entire growing season. In conclusion, N nutrition in the studied forest ecosystem seems to be optimally attuned to European beech.     

The impact of nitrogen deposition on carbon sequestration by European forests and heathlands

In this study, we present estimated ranges in carbon (C) sequestration per kg nitrogen (N) addition in above-ground biomass and in soil organic matter for forests and heathlands, based on: (i) empirical relations between spatial patterns of carbon uptake and influencing environmental factors including nitrogen deposition (forests only), (ii) ¹⁵N field experiments, (iii) long-term low-dose N fertilizer experiments and (iv) results from ecosystem models. The results of the various studies are in close agreement and show that above-ground accumulation of carbon in forests is generally within the range 15–40 kg C/kg N. For heathlands, a range of 5–15 kg C/kg N has been observed based on low-dose N fertilizer experiments. The uncertainty in C sequestration per kg N addition in soils is larger than for above-ground biomass and varies on average between 5 and 35 kg C/kg N for both forests and heathlands. All together these data indicate a total carbon sequestration range of 5–75 kg C/kg N deposition for forest and heathlands, with a most common range of 20–40 kg C/kg N. Results cannot be extrapolated to systems with very high N inputs, nor to other ecosystems, such as peatlands, where the impact of N is much more variable, and may range from C sequestration to C losses.     

Impact of water scarcity on spruce and beech forests

One of the greatest threats posed by ongoing climate change may be regarded the drought caused by changes in precipitation distribution. The aim of presented study was to characterize reactions to dry conditions and conditions without drought stress on gross primary production (GPP) and net ecosystem production (NEP) of spruce and beech forests, as these two species dominate within the European continent. Daily courses of GPP and NEP of these two species were evaluated in relation to an expected decrease in CO2 uptake during dry days. The occurrence of CO2 uptake hysteresis in daily production was also investigated. Our study was performed at Bily Kriz(spruce) and Stitna(beech) mountain forest sites during 2010–2012 period. We applied eddy covariance technique for the estimation of carbon fluxes, vapor pressure deficit and precipitation characteristics together with the SoilClim model for the determination of drought conditions, and the inverse of the Penman–Monteith equation to compute canopy conductance. Significant differences were found in response to reduced water supply for both species. Spruce reacts by closing its stomata before noon and maintaining a reduced photosynthetic activity for the rest of the day, while beech keeps its stomata open as long as possible and slightly reduces photosynthetic activity evenly throughout the entire day. In the spruce forest, we found substantial hysteresis in the light response curve of GPP. In the beech forest, the shape of this curve was different: evening values exceeded morning values.     

Soil functional indicators in mixed beech forests are clearly species-specific

Beech stands are considered part of the ancient forest ecosystems in the northern hemisphere.In mixed stands in beach forest ecosystems,the type of associated tree species can significantly affect soil functions,but their influence on microbial activity,nutrient cycling and belowground properties is unknown.Here,we considered forest patches in northern Iran that are dominated by different tree species:Fagus orientalis Lip sky,Quercus castaneifolia C.A.Mey.,Pterocarya fraxinifolia(Lam.),Tilia beg...     

Effect of canopy gap size and ecological factors on species diversity and beech seedlings in managed beech stands in Hyrcanian forests

We studied the species diversity of the herb layer and ecological factors in harvest-created gaps in beech stands under a single-tree selection system in Northern Iran. To determine diversity, the number of beech seedlings, and other ecological factors, 16 gaps were selected and subplots of 5 m2 were positioned at the centre and at the cardinal points of each gap. Species richness and Simpson diversity index increased with increasing gap area as did numbers of seedlings. With increasing humus layer thickness, species richness declined but the Hill evenness index increased. Species richness increased with increasing light availability. There was no relationship between crown radii of beech trees and diversity indices. Correlations between environmental factors and numbers ofindividuals of some species in the herb layer were not significant except in a few cases. The results help explain the effects of man-made gaps on the dynamics of managed beech stands and this benefits evaluation of silvicultural operating plans.     

Long-term trends in vegetation changes of managed versus unmanaged Eastern Carpathian beech forests

A vegetation survey of semi-permanent plots was conducted between years 1955–1962 and 2005–2009, and aimed to determine floristic changes occurring in beech forests located on a scarcely populated, dense forest area. The survey encompassed all natural beech forest types within the Polish part of the “Eastern Carpathians” International Reserve of Biosphere. A comparison of trends in changes between unmanaged (Bieszczady National Park) and managed (the remaining part of the Biosphere Reserve) forests was attempted. DCA analysis was used for the determination of changes on the community level. The density of vegetation, species richness, frequency of occurrence and cover, and functional groups of species were determined. Much more profound community changes were determined to have occurred in managed beech forests. However, a similar direction in vegetation changes within both managed and unmanaged forests verifies the existence of a regional pattern of changes in beech forests. This regional pattern involves (1) a decrease in the shares of arborescent species within the shrub layer, (2) disturbance of the canopy layer, (3) acidification of the top layer of the soil, (4) change of light conditions, (5) increase in shares of generalist species and decrease in shares of specialist species. A decreasing anthropogenic pressure, aging of forest stands, functioning of large, dense forest areas and sustainable forest management strategies forestry management system mimicking natural deciduous forest disturbance regime constitute factors shaping the regional changes of the forest vegetation. The undergoing changes can lead to biotic and spatial homogenization of the Eastern Carpathian beech forests.     

Water-holding characteristics of litter in different forests at the Lianxiahe watershed

We surveyed the forest litter amount at the Lianxiahe watershed in the Three Gorges Reservoir Area and analyzed some hydrological characteristics such as the maximum water-holding capacity and water absorption rate of litter in six types of forests, i.e. the Cupressus funebris forest, the coniferous mixed forest, the coniferous and broad-leaved mixed forest, the broad-leaved forest, the Pinus massoniana forest, the bush forest. Results showed that the litter amount follows the order of the coniferous and broad-leaved mixed forest > the pure C. funebris forest, the P. massoniana forest > the coniferous mixed forest > the broad-leaved forest > the bush forest. The maximal water holding capacity of the undecomposed litter is in the order of the C. funebris forest > the coniferous mixed forest > the bush forest > the coniferous and broad-leaved mixed forest > the broad-leaved forest > the P. massoniana forest. The maximal water-holding capacity of the half-decomposed litter is the P. massoniana forest > the coniferous and broad-leaved mixed forest > the C. funebris forest > the coniferous mixed forest > the broad-leaved forest > the bush forest. In this watershed, the water holding capacity of the litter in the C. funebris forest is the highest, followed by the coniferous mixed forest, coniferous and broad-leaved mixed forest, P. massoniana forest, broad-leaved forest and bush forest. When the soil is covered only by litter, both the maximal rainfall amount and intensity in different forest stands are different if there is no water infiltration and runoff from the ground surface. __________ Translated from Journal of Huazhong Agricultural University, 2005, 27(2): 207–212 [译自: 华中农业大学学报, 2005, 24(2): 207–212]     

The north-eastern distribution range of European beech a review

Today, European beech (Fagus sylvatica L.) seems to be a markedlysuccessful tree species in the north-east of its distributionrange. The distribution area may be larger than originally assumed;past forest management is probably the main cause of the contractionin the postglacial European beech range. Numerous attempts consistentlyhave failed to locate a distinct distribution edge for Europeanbeech. Therefore, we define northern and eastern Poland andthe southern regions of the Baltic States as margins of Europeanbeech distribution. Ecophysiological approaches have identifiedthe drought constraints for European beech in terms of (1) thecritical limit for xylem cavitation and loss of hydraulic conductivity,reached at a shoot water potential of –1.9 MPa, and (2)a reduction in gross primary production and total ecosystemrespiration when relative extractable soil water reaches 40and 20 per cent, respectively. However, it is difficult to correlateEuropean beech distribution margins with single macro-climaticfactors. Moreover, the adaptation of European beech populationsand provenances to drought and frost varies. The phenotypicplasticity and evolutionary adaptability of European beech appearto be underestimated. These characteristics may counteract afurther contraction of the European beech range arising fromclimate change in the future.     

Disturbances in European beech water relation during an extreme drought

Context

In the context of a probable increase in intensity and frequency of extreme summer drought events, a better understanding of the key processes involved in water relations is needed to improve the theoretical foundations of predictive process-based models.

Aims

This paper aims to analyse how temperate deciduous trees cope with water shortage.

Methods

The exceptional summer drought of 2003 in Europe provided an opportunity to monitor stomatal conductance and twig water potential in European beech (Fagus sylvatica L.) at predawn and midday and to analyse variations with respect to leaf height within the canopy. By comparing our field measurements of twig water potential to values found in the literature, we confirmed the strong impact of soil water shortage on crown water relations.

Results

This paper shows that (1) the vertical gradient of stomatal conductance within the crown disappeared under extreme soil water depletion; (2) at maximum drought intensity, predawn twig water potential (ψ _pd) reached ?2.3 MPa at a height of 14 m in the crown and ?2.0 MPa at a height of 10 m. The significant differences in ψ _pd between the two measurement heights in the canopy may be due to night transpiration; (3) there was a close relationship between predawn twig water potential and relative extractable soil water; (4) as drought conditions intensified, there was a close relationship between canopy radiation interception and predawn water potential, as estimated daily from relative extractable soil water.