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.     

Climate influences the leaf area/sapwood area ratio in Scots pine

We tested the hypothesis that the leaf area/sapwood area ratio in Scots pine (Pinus sylvestris L.) is influenced by site differences in water vapor pressure deficit of the air (D). Two stands of the same provenance were selected, one in western Scotland and one in eastern England, so that effects resulting from age, genetic variability, density and fertility were minimized. Compared with the Scots pine trees at the cooler and wetter site in Scotland, the trees at the warmer and drier site in England produced less leaf area per unit of conducting sapwood area both at a stem height of 1.3 m and at the base of the live crown, whereas stem permeability was similar at both sites. Also, trees at the drier site had less leaf area per unit branch cross-sectional area at the branch base than trees at the wetter site. For each site, the average values for leaf area, sapwood area and permeability were used, together with values of transpiration rates at different D, to calculate average stem water potential gradients. Changes in the leaf area/sapwood area ratio acted to maintain a similar water potential gradient in the stems of trees at both sites despite climatic differences between the sites.     

Response of LAI-2000 estimates to changes in plant surface area index in a Scots pine stand

We assessed the accuracy with which the LAI-2000 plant canopy analyzer measured changes in leaf area index (LAI) and plant area index (PAI) in a 25-year-old Scots pine (Pinus sylvestris L.) stand. Stand density was 2100 stems ha(-1) and mean tree height was 8.7 m. Needle and branch areas of the stand were reduced progressively to zero by the stepwise removal of branches on all trees growing in a circular plot with a radius of 25 m. An LAI-2000 estimate was taken after each step reduction. The needle and branch surface areas removed at each step were estimated from direct measurements and were compared with the changes in the LAI-2000 estimates. Initially (before removal of branches), directly measured PAI was 5.2 (needles = 86%, branches = 8% and stems = 6%). The LAI-2000 estimate of total surface area was 66% of direct PAI and 77% of direct LAI. There was a nonlinear relationship between the LAI-2000 estimate and directly measured PAI, such that their ratio (equivalent to the clumping factor) increased from 0.66 to 1.05 with decreasing PAI. At the last measurement, when only stems were left, the LAI-2000 estimate agreed well with the direct measurement of PAI. The LAI-2000 underestimated the direct measurement of LAI at the first three steps when LAI was > 2 and the proportion of woody area was small (< 20%). However, because the LAI-2000 estimate included stem and branch areas, it overestimated the direct measurement of LAI at the last three measurements when the proportion of woody area was large (> 20%).     

Tree effects on the chemical topsoil features of oak, beech and pine forests

We aimed to study tree effects on the chemical properties of forest soils. We compared soil features of three types of forest ecosystems, each with four stands (replicates): beech forests (Fagus sylvatica), oak forests (dominated by Quercus pyrenaica) and pine plantations (Pinus sylvestris). Five samples from the top 10 cm of soil were taken per stand, from which pH, organic matter content (O.M.), total nitrogen (N) and available calcium (Ca²⁺), magnesium (Mg²⁺), potassium (K⁺) and sodium (Na⁺) were determined. Litter layer depth was measured at each soil sampling point. We also measured tree density and crown diameters at each stand. Our results indicated that soil samples from the four pine plantation stands were more similar while oak and beech stands were characterised by great variability in terms of soil properties and leaf litter depth. Although the identity of the dominant tree species significantly influenced several topsoil chemical properties (increase in pH and available cations in oak forests and higher organic matter and total nitrogen in beech and pine ecosystems), there were other important factors affecting soil features that may be taken under consideration. Differences between soil properties of the three types of forest ecosystems were mainly related to the characteristics of the litter layer and less related to the tree layer structure. Finally, the establishment of pine plantations in naturally deciduous tree areas made the topsoil features more homogeneous.     

Comparing the influence of site quality,stand age,fire and climate on aboveground tree production in Siberian Scots pine forests

Temporal patterns of stem and needle production and total aboveground net primary production (ANPP) were studied at the tree and stand level along four chronosequences of Siberian Scots pine (Pinus sylvestris L.) forests differing in site quality (poor lichen type or the more fertile Vaccinium type) and in frequency of surface fires (unburned, moderately burned (fire return interval of approximately 40 years), or heavily burned (fire return interval of approximately 25 years)). The maximum range of variability in aboveground production was quantified for: (1) possible long-term changes in site quality; (2) stand age; (3) non-stand-replacing, recurring surface fires; and (4) interannual climate variability. For (1) and (2), total ANPP was low in the lichen-type chronosequence, reached a maximum of 170 g C m(-2) year(-1) after 100 years and decreased to 100 g C m(-2) year(-1) in older stands. Maximum ANPP in the Vaccinium-type chronosequence was 340 g C m(-2) year(-1) and occurred earlier in the 53-year-old stand than in the other stands. Along the lichen-type chronosequences, peak ANPP was paralleled by maximum carbon allocation to stem growth. (3) In mature trees, damage by recurrent surface fires decreased stem growth by 17 +/- 19% over a 10-year period relative to pre-fire values. At longer timescales, ANPP was hardly affected by fire-related differences in mortality. (4) Needle- plus stem-NPP, reconstructed for a 3-year period, varied within a range of 15 g C m(-2) year(-1) in the lichen-type stands and 35 g C m(-2) year(-1) in the Vaccinium-type stands. For the same period, the coefficient of variance was higher for needle-NPP (20 +/- 10%) than for stem-NPP (12 +/- 7%). Needle- and stem-NPP did not covary in time. Most 30-year time series of stem-NPP at the tree level exhibited strong autocorrelation. In older trees, stem-NPP was positively correlated with growing season precipitation. Thus, the factors driving variability in ANPP ranked according to their maximum influence as: stand age (controlled by the frequency of stand-replacing fires) > site quality > growth depression because of surface fire damage approximately equal age-related reduction in ANPP > interannual variability approximately equal long-term effects of fire (stand density reduction). In lichen-type forests, we found that ANPP at the landscape level declined sharply when the interval between stand-replacing fires was less than 120 years, illustrating that fire strongly influences ANPP of boreal Scots pine forests.     

冀北山地油松林净生产力与相关立地因子关系模型的研究

为推测林分生产力,提高森林经营水平,在对冀北山地油松林标准地进行调查的基础上,建立树高随林龄的增长而变化的导向曲线模型,通过模型给出各标准地的立地指数;建立油松单株生物量与单株胸径树高关系模型,导出各标准地林分生物量;建立林分生物量与林分密度、林龄及立地指数关系模型,进而建立林分净生产力与林分密度、林龄及立地指数关系模型。     

Fine root biomass in relation to site and stand characteristics in Norway spruce and Scots pine stands

Variations in fine root biomass of trees and understory in 16 stands throughout Finland were examined and relationships to site and stand characteristics determined. Norway spruce fine root biomass varied between 184 and 370 g m(-2), and that of Scots pine ranged between 149 and 386 g m(-2). In northern Finland, understory roots and rhizomes (< 2 mm diameter) accounted for up to 50% of the stand total fine root biomass. Therefore, the fine root biomass of trees plus understory was larger in northern Finland in stands of both tree species, resulting in a negative relationship between fine root biomass and the temperature sum and a positive relationship between fine root biomass and the carbon:nitrogen ratio of the soil organic layer. The foliage:fine root ratio varied between 2.1 and 6.4 for Norway spruce and between 0.8 and 2.2 for Scots pine. The ratio decreased for both Norway spruce and Scots pine from south to north, as well as from fertile to more infertile site types. The foliage:fine root ratio of Norway spruce was related to basal area and stem surface area. The strong positive correlations of these three parameters with fine root nitrogen concentration implies that more fine roots are needed to maintain a certain amount of foliage when nutrient availability is low. No significant relationships were found between stand parameters and fine root biomass at the stand level, but the relationships considerably improved when both fine root biomass and stand parameters were calculated for the mean tree in the stand. When the northern and southern sites were analyzed separately, fine root biomass per tree of both species was significantly correlated with basal area and stem surface area per tree. Basal area, stem surface area and stand density can be estimated accurately and easily. Thus, our results may have value in predicting fine root biomass at the tree and stand level in boreal Norway spruce and Scots pine forests.     

Effects of climate and site characteristics on Scots pine growth

Scots pine is a highly diverse species, extended across Europe from Scandinavia to Spain, Italy, Greece and Turkey. It is also a valuable species, used in many commercial monoculture plantations in Great Britain and particularly in Scotland. Because of the diversity of growing environments and its commercial importance, it is necessary to identify the combination of significant factors affecting the observed variability of growth. Temperature, mainly during the growing season, is quite commonly considered as the most important factor in knowledge-based or empirical models. However, in highly oceanic climates like that of Scotland, the impact of temperature may have a less significant impact on growth. Here we argue that other factors, such as incoming winter solar radiation, frost, drought and management also have a significant effect on the growth of Scots pine. In addition, we argue that the already developed Ecological Site Classification knowledge-based model, used as a forest management tool in Great Britain, should be updated to incorporate our findings. Furthermore, we discuss the need to include management impact and possibly more physiological based components in its growth modelling routines, as these would allow the introduction of the effect of winter solar radiation.     

Scots pine stand establishment with special emphasis on uncertainty and cost-effectiveness,the case of northern Finland

The effect of uncertainty on the cost-effectiveness of alternative chains of stand establishment in northern Finland was examined. The data were from a reforestation study of Scots pine (Pinus sylvestris L.) consisting of 288 sample plots, which were measured with respect to regeneration success. The study design included four site-preparation methods (patch scarification, ploughing, prescribed burning and disk trenching) combined with three reforestation methods (sowing, planting with containerized seedlings and planting with bare-rooted transplants). Initial reforestation density was 2,500 spots or seedlings per hectare, and the regeneration success was modeled as probability with two thresholds, namely 500 and 1,100 saplings. On formerly spruce-dominated as well as pine-dominated sites the most cost-effective chain was ploughing and planting with containerized seedlings, when threshold was 1,100 saplings per hectare. However, with threshold of 500 saplings the best performer was ploughing and direct sowing on both sites.     

人工红松幼龄林的叶面积指数与生长动态

对凉山自然保护区的全坡位红松（Ｐｉｎｕｓ ｋｏｒａｉｅｎｓｉｓ）造林地进行了生长与叶面积指数的动态调查表明，立地条件好的地段红松生长好，上层阔叶树发育亦好。立地条件和上层阔叶树的庇护是导致红松生长差异的主要原因。     

Measurements of fungal wood decay on Scots pine and beech by means of X-ray microdensitometry

In this work, we present a fast and promising method to evaluate the natural durability of wood based on X-ray microdensitometry. Tested on beech and Scots pine wood samples, our findings show that this methodology and the traditional EN standards methodology based on mass loss are strongly correlated. X-ray methodology is less time consuming (we can detect the effectiveness of the attack within 5–6 weeks) and less expensive (very cheap plastic Petri dishes instead of the expensive glass Kolle flasks); moreover, the proposed method allows to thoroughly examine the phases and the kinetics of the fungal attack, and to investigate the spatial repartition of the attack within the samples due to the low thickness of the sample.     

Comparison of soil nitrogen dynamics under beech,Norway spruce and Scots pine in central Germany

To investigate the effect of tree species on soil N dynamics in temperate forest ecosystems, total N (Nt), microbial N (Nmic), net N mineralization, net nitrification, and other soil chemical properties were comparatively examined in beech (64–68 years old) and Norway spruce (53–55 years old) on sites 1 and 2, and beech and Scots pine (45 years old) on site 3. The initial soil conditions of the two corresponding stands at each site were similar; soil types were dystric Planosol (site 1), stagnic Gleysols (site 2), and Podzols (site 3). In organic layers (LOf₁, Of₂, Oh), Nmic and Nmic/Nt, averaged over three sampling times (Aug., Nov., Apr.), were higher under the beech stands than under the corresponding coniferous ones. However, the Nmic in the organic layers under beech had a greater temporal variation. Incubation (10 weeks, 22 °C, samples from November) results showed that the net N mineralization rates in organic layers were relatively high with values of 8.1 to 24.8 mg N kg^–1 d^–1. Between the two corresponding stands, the differences in net N mineralization rates in most of the organic layers were very small. In contrast, initial net nitrification rates (0.2–17.1 mg N kg^–1 day^–1) were considerably lower in most of the organic layers under the conifer than under the beech. In the mineral soil (0–10 cm), Nmic values ranged from 4.1–72.7 mg kg^–1, following a clear sequence: August>November>April. Nmic values under the beech stands were significantly higher than those under the corresponding coniferous stands for samples from August and April, but not from November. The net N mineralization rates were very low in all the mineral soils studied (0.05–0.33 mg N kg^–1 day^–1), and no significant difference appeared between the two contrasting tree species.     

Temporal variation and efficiency of leaf area index in young mountain Norway spruce stand

Temporal variation of leaf area index (LAI) in two young Norway spruce stands with different densities was monitored during eight consecutive growing seasons (1998–2005). We focused on: (1) LAI dynamics and above-ground mass production of both spruce stands and their comparison, (2) leaf area duration (LADU), crop production index (CPI) and leaf area efficiency (LAE) evaluation, and (3) thinning impact on the above-mentioned parameters. Also, we tried to deduce the most effective LAI value for the Norway spruce forest investigated. The LAI values of both spruce stands showed a typical seasonal course. To describe the LAI dynamics of the stand, we recommend taking LAI measurements within short time intervals at the time of budding and needle expansion growth (i.e., in early spring) and close to the LAI peak, when the twig growth has been completed. The reason was that after reaching the seasonal maximum, no significant differences between subsequently obtained values were found in the following 2 months. Therefore, we recommend this period for the estimation of seasonally representative LAI values, enabling the comparison of various spruce stands. The maximum hemi-surface LAI value reached 12.4. Based on our results, the most effective LAI values for maximum above-ground biomass production were within the range of 10–11. We found an LAI over these values to be less effective for additional production of above-ground biomass. In forest practice, thinning intensity is mostly described by percentage of stocking reduction. We want to show that not only thinning intensity, but also the type of thinning is important information. The type of thinning significantly affected the stand above-ground biomass increment, canopy openness, stand LAI and LAI efficiency. The stimulating effect of high-type thinning was observed; the LAE as well as the CPI increased. Low-type thinning had no such effects on LAE increments compared to the high-type thinning with similar intensity.

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Persisting soil drought reduces leaf specific conductivity in Scots pine (Pinus sylvestris) and pubescent oak (Quercus pubescens)

Leaf specific conductivity (LSC; the ratio of stem conductivity (K(P)) to leaf area (A(L))), a measure of the hydraulic capacity of the stem to supply leaves with water, varies with soil water content. Empirical evidence for LSC responses to drought is ambiguous, because previously published results were subject to many confounding factors. We tested how LSC of similar-sized trees of the same population, under similar climatic conditions, responds to persistently wet or dry soil. Scots pine (Pinus sylvestris L.) and pubescent oak (Quercus pubescens Willd.) trees were compared between a dry site and a wet site in the Valais, an inner alpine valley in Switzerland. Soil water strongly influenced A(L) and K(P) and the plant components affecting K(P), such as conduit radius, conduit density and functional sapwood area. Trees at the dry site had lower LSC than trees with the same stem diameter at the wet site. Low LSC in trees at the dry site was associated with a smaller functional sapwood area and narrower conduits, resulting in a stronger reduction in K(P) than in A(L). These observations support the hypothesis that trees maintain a homeostatic water pressure gradient. An alternative hypothesis is that relatively high investments in leaves compared with sapwood contribute to carbon gain over an entire season by enabling rapid whole-plant photosynthesis during periods of high water availability (e.g., in spring, after rain events and during morning hours when leaf-to-air vapor pressure deficit is small). Dynamic data and a hydraulic plant growth model are needed to test how investments in leaves versus sapwood and roots contribute to transpiration and to maximizing carbon gain throughout entire growth seasons.     

Trajectory analysis of long-term changes in the nutritional status of a Scots pine stand

For the Scots pine (Pinus sylvestris L.) stand Pustert (Bavaria, Germany), the temporal change in its nutritional status as assessed by foliar analysis is analyzed using different techniques of statistical data evaluation and graphical representation. The procedures include new methods such as the development of adequate trajectories and their combination with an improved “Timmer and Stones graphical diagnostic system” as well as with a Principal Component Analysis (PCA). This mothod of data evaluation and presentation has several advantages: (i) It allows to visualize the dynamics of a multi-dimensional data set in two-dimensional graphs. Additionally, (ii) it provides a tool to easily identify various types of temporal changes in tree nutrition as e.g. nutrient insufficiency, luxury consumption, and dilution. Finally (iii), it enables the reader to distinguish quickly between non-systematic inter-annual variation, episodic cyclic changes, and systematic trends in the nutritional status of a forest stand. For the studied site, three major changes in the nutritional status of the Scots pine stand during the past 30 years could be identified: (i) a concomitant improvement of its N and P nutrition between 1964 and 1980 was observed, primarily reflecting the recovery of the ecosystem from past intensive litter-raking and other exploitative forest utilization. We also recognized (ii) a trend towards an unbalanced nutrient supply induced by excessive gains of the ecosystem in N compared to other major nutrients particularly after 1980, which is probably due to elevated atmospheric N deposition. In particular, we noticed (iii) a continuous dilution of the Ca concentrations in the foliage. Obviously, Ca uptake could not keep pace with the increased biomass accumulation. Dedicated to Professor Dr.K. E. Rehfuess to his 65^th birthday     

Identifying dendroecological growth releases in American beech,jack pine,and white oak: Within-tree sampling strategy

The objective of this project was to identify the timing of growth release events detected from tree ring widths and compare whether two cores taken from the same tree reconstructed the same disturbance history. This research question is important because current dendroecological reconstructions of canopy disturbance rely on sampling one core per tree; however, the variation of releases from different cores from the same tree has never been evaluated. We sampled two increment cores from 20 jack pine, 17 white oak, and 19 American beech and identified release events with two commonly employed methods: radial growth averaging technique and boundary line criteria. In jack pine, 85% of the paired cores showed identical releases with the radial growth averaging technique, but 15% of the paired cores varied in reconstructed growth releases. In the jack pine, no releases were identified with the boundary line criteria for any of the paired cores. In the white oak, 65% had identical releases identified with the radial growth averaging technique and 35% of the pairs showed differences. The boundary line criteria for white oak had agreement between releases for 76% of the pairs and different release histories for 24% of the pairs. In the American beech, we were only able to use the radial growth averaging technique and this method showed identical release timing for 79% of the paired cores and differences in 21% of the paired cores. This level of within-tree growth variation is unlikely to influence identification of stand-wide disturbances; however, for reconstructions of small-scale disturbances it is likely to under-represent disturbance events. Therefore, for small-scale disturbance reconstructions, we recommend dendroecologists consider sampling two cores per tree instead of the standard sample of one core per tree.     

Regional influences of soil available water-holding capacity and climate,and leaf area index on simulated loblolly pine productivity

We simulated loblolly pine (Pinus taeda L.) net canopy assimilation, using BIOMASS version 13.0, for the southeastern United States (1° latitude by 1° longitude grid cells) using a 44-year historical climate record, estimates of available water-holding capacity from a natural resource conservation soils database, and two contrasting leaf area indices (LAI) (low; peak LAI of 1.5 m² m⁻² projected, and high; 3.5 m² m⁻²). Median (50th percentile) available water-holding capacity varied from 100 to 250 mm across the forest type for a normalized 1.25 m soil profile. Climate also varied considerably (growing season precipitation ranged from 200 to 1600 mm while mean growing season temperature ranged from 13° to 26°C). Net canopy assimilation ranged from 9.3 to 19.2 Mg C ha⁻¹ a⁻¹ for high LAI and the 95th percentile of available water-holding capacity simulations.We examined the influence of soil available water-holding capacity, and annual variation in temperature and precipitation, on net canopy assimilation for three cells of similar latitude. An asymptotic, hyperbolic relationship was found between the 44-year average net canopy assimilation and soil available water-holding capacity. Shallow soils had, naturally, low water-holding capacity (<100 mm) and, subsequently, low productivity. However, median available water-holding capacity (125–150 mm) was sufficient to maintain near maximum production potential in these cells.Simulations were also conduced to examine the direct affects of soil available water on photosynthesis (P_N) and stomatal conductance (g_S) on net canopy assimilation. In the absence of water limitations on P_N and g_S, net canopy assimilation increased by only 10% or less over most of the loblolly pine region (when compared to simulations for median available water-holding capacity with water influences in place). However, the production differences between high and low LAI, at the median soil available water-holding capacity, ranged from 30% to 60% across the loblolly pine range. Vapor pressure deficit was found to dramatically reduce productivity for stands of similar LAI, incident radiation, rainfall, and available water-holding capacity. Thus, these simulations suggest that, regionally, loblolly pine productivity may be more limited by low LAI than by soil available water-holding capacity (for soils of median available water-holding capacity or greater). In addition, high atmospheric forcing for water vapor will reduce net assimilation for regions of otherwise favorable available water and LAI.     

Influence of nurse birch and Scots pine seedlings on early aerial development of European beech seedlings in an open-field plantation of Central France

Beech woodland can be restored by direct planting of beech (Fagussylvatica L.) seedlings in abandoned areas, but this methodis generally avoided in forestry because of the growth difficultiesof beech in full-light conditions. This study tested a methodthat consists of planting beech seedlings in full-light conditionswith silver birch (Betula pendula Roth.) or Scots pine (Pinussylvestris L.) as nurse trees. A total of 65 two-year-old beechseedlings surrounded by either 3, 4, 5 or 6 pine or birch seedlingsor without competitors were planted in bare-soil open-fieldconditions in Central France. Tree growth and light availabilitywere monitored at the end of each of the following 3 years,and soil moisture was also measured the last year. At the endof the experiment, aerial biomass measurements were made onthe beeches. Results showed that relative beech growth in termsof diameter was significantly reduced by the local neighbouringtrees and that this reduction was particularly noticeable inthe pine treatments where there was extensive pine seedlingdevelopment. In contrast, relative beech height did not exhibitany significant variations among the treatments. Light availabilitywas decreased by the neighbouring trees, especially in the pinetreatments. Light reduction was more pronounced in the last2 years and in the middle or lower parts of the canopy of thesubject beech. Soil water content was lower under the pine canopiesthan under the birch canopies, and was positively correlatedto beech relative diameter growth. Specific leaf area, height-to-diameterratio and crown length-to-crown width ratio weakly but significantlyincreased with competition from the neighbours. Beech biomasspartitioning was only weakly affected by the treatments. Thisstudy showed that nurse trees tend to reduce beech growth butimprove form although effects remained weak due to the insufficientlength of the experiment. Further studies are also needed toquantify the effects of the neighbourhood on the growth of competitiveherbaceous vegetation and on changes in the microclimatic conditions.