Potential productivity of forested areas based on a biophysical model. A case study of a mountainous region in northern Spain

? Today’s forest managers face a number of important challenges involving an increasing need for precise estimates of forest structure and biomass, potential productivity or forest growth. The objective is to develop a model for potential productivity in a mountainous region of Spain. The model combines climatic, topographic and lithological data using a variant of a traditional biophysical model: the Paterson index.

? In a first approach, the climatic productivity is assessed by modelling the required parameters using different geostatistical techniques and software supported by GIS. A second approach includes the correction of the former productivity classes considering the different lithological facies. The potential forest productivity model involves the integration of both models.

? Finally, data from the National Forest Inventory (NFI) are used to compare the real and potential yield data within different regions of the studied area.

? The results of these analyses demonstrate the usefulness of the model, particularly in mountainous regions, where no significant differences are found between the data from the NFI and the model, but they also show the discrepancies between the estimates and real data when the latter are considered for different tree species, diameter classes or management.

     

Ontogenetic changes in stomatal and biochemical limitations to photosynthesis of two co-occurring Mediterranean oaks differing in leaf life span

A quantitative analysis was applied to the stomatal and biochemical limitations to light-saturated net photosynthesis under optimal field conditions in mature trees and seedlings of the co-occurring evergreen oak, Quercus ilex L., and the deciduous oak, Q. faginea Lam. Stomatal limitation to photosynthesis, maximal Rubisco activity and electron transport rate were determined from assimilation versus intercellular leaf carbon dioxide concentration response curves of leaves that were subsequently analyzed for nitrogen (N) concentration, mass per unit area, thickness and percent internal air space. In both species, seedlings had a lower leaf mass per unit area, thickness and leaf N concentration than mature trees. The root system of seedlings during their third year after planting was dominated by a taproot. A lower leaf N concentration of seedlings was associated with lower maximal Rubisco activity and electron transport rate and with assimilation rates similar to or lower than those of mature trees, despite the higher stomatal conductances and potential photosynthetic nitrogen-use efficiencies of seedlings. Consequently, stomatal limitation to photosynthesis increased with tree age in both species. In both seedlings and mature trees, a lower assimilation rate in Q. ilex than in Q. faginea was associated with lower stomatal conductance, N allocation to photosynthetic functions, maximal Rubisco activity and electron transport rate, and potential photosynthetic nitrogen-use efficiency but greater leaf thickness and leaf mass per unit area. Tree-age-related changes differed quantitatively between species, and the characteristics of the two species were more similar in seedlings than in mature trees. Despite higher stomatal conductances, seedlings are more N limited than adult trees, which contributes to lower biochemical efficiency.     

Flux partitioning in an old-growth forest: seasonal and interannual dynamics

Turbulent fluxes of carbon, water and energy were measured at the Wind River Canopy Crane, Washington, USA from 1999 to 2004 with eddy-covariance instrumentation above (67 m) and below (2.5 m) the forest canopy. Here we present the decomposition of net ecosystem exchange of carbon (NEE) into gross primary productivity (GPP), ecosystem respiration (R(eco)) and tree canopy net CO(2) exchange (DeltaC) for an old-growth Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco)-western hemlock (Tsuga heterophylla (Raf.) Sarg.) forest. Significant amounts of carbon were recycled within the canopy because carbon flux measured at the below-canopy level was always upward. Maximum fluxes reached 4-6 micromol m(-2) s(-1) of CO(2) into the canopy air space during the summer months, often equaling the net downward fluxes measured at the above-canopy level. Ecosystem respiration rates deviated from the expected exponential relationship with temperature during the summer months. An empirical ecosystem stress term was derived from soil water content and understory flux data and was added to the R(eco) model to account for attenuated respiration during the summer drought. This attenuation term was not needed in 1999, a wet La Ni?a year. Years in which climate approximated the historical mean, were within the normal range in both NEE and R(eco), but enhanced or suppressed R(eco) had a significant influence on the carbon balance of the entire stand. In years with low respiration the forest acts as a strong carbon sink (-217 g C m(-2) year(-1)), whereas years in which respiration is high can turn the ecosystem into a weak to moderate carbon source (+100 g C m(-2) year(-1)).     

Ranking temperate woody species along a gradient of browsing by deer

As a result of diverse management policies, densities of red and roe deer have varied considerably for the last few decades in Europe. Their selective browsing on woody species can modify the interactions between competing species and thus change overall patterns of plant diversity in forests. However, no classification of woody species according to browsing by deer yet exists in Europe. The selectivity index, which measures the balance between availability and consumption of plant species by herbivores, could be used for such a classification. The aim of this study was, first, to rank woody species according to this selectivity index, then to test whether their rank changed when browsing pressure varied in space or time. We also explored morphological and ecological life traits (Ellenberg indicator values) that might explain differences in selectivity.In 2006, we re-sampled a network of 217 coupled floristic and browsing survey plots, which had been previously described twice (1976, 1981). The plots were located in the forest of Arc-en-Barrois (France), a large woodland area divided into a North and a South Forest which have had different histories of browsing pressure.For the three samples and the two forests, we calculated a selectivity index for 19 woody species. Dogwood (Cornus sp.) and field rose (Rosa arvensis) appeared to be the most selected species whereas beech (Fagus sylvatica), common mezereon (Daphne mezereum) and wild-service tree (Sorbus torminalis) were always avoided. The rank order of species according to their selectivity index remained stable over the three sample years and between the two forests (Spearman's rank correlations: ρ ≈ 0.8 spatially, and range from 0.3 to 0.8 temporally). This indicates that variations in browsing pressure did not strongly modify deer selectivity patterns. Selectivity was positively correlated with wood density and negatively correlated with maximum height of the species, but unrelated to other plant traits. Finally, selectivity can be seen as an intrinsic characteristic of those woody species, and could be used as an additional life trait when analyzing vegetation communities.     

Forest monitoring with TerraSAR-X: first results

Several TerraSAR-X satellite images acquired in high resolution spotlight mode with different polarisations for two study sites in southern Germany were used to distinguish forest from other land cover classes (agriculture, built-up, water bodies) using logistic regression models. In general, we observed that the mean and particularly the standard deviation of the backscatter were viable measures to discriminate land cover classes. Both measures were lowest for water bodies and highest for built-up areas, with agricultural areas and forest in intermediate positions. Trees outside forest were not differentiable from forest with the applied method. The HH-polarised image was better suited for a classification of built-up areas, whereas the VV-polarised image was more appropriate when classifying agricultural areas. Consequently, the combination of the two differently polarised images leads to a significantly better model. Since forests in one study area were generally found on steeper slopes in comparison to other land cover classes, the inclusion of terrain slope further improved the classification, which resulted in an overall accuracy of 92–95%. Systematic differences in the parameter values of the explanatory variables for one class between the study areas may be caused by differing class probabilities. Thus, transferring the model of one study area to the image of another resulted in a 7–9% loss of accuracy.     

Impact of deer on temperate forest vegetation and woody debris as protection of forest regeneration against browsing

A 3-year field experiment with paired exclosure (fenced areas, excluding deer) and control plots (unfenced areas, free access to deer), with two treatments with and without woody debris, was carried out at two sites in a temperate forest in eastern France. The aim of the experiment was to assess the effect of browsing by roe deer (Capreolus capreolus) and red deer (Cervus elaphus) on the diversity and richness of plant species and to test the effectiveness of using woody debris to protect seedlings and saplings from deer browsing. In presence of deer, both plant species richness and diversity were reduced the first year of the study, but this negative impact of deer then disappeared after 3 years. Deer browsing mostly affected species composition of plant communities. We observed a decrease in the abundance of preferred species such as Carpinus betulus, Rubus fructicosus, Rubus idaeus, Anemone nemorosa and Epilobium angustifolium, and palatable species such as Acer spp., Carex spp., Festuca spp. and Mycelis muralis, whereas unpalatable species such as Lamium spp., or species particularly resilient to browsing such as grasses (Brachypodium spp. and Luzula spp.) increased in abundance. The use of woody debris as protection against browsing by deer did not limit damage to seedlings and saplings of the main commercially valuable species, Abies alba and Quercus spp. Instead of limiting deer impact, use of woody debris seemed to increase the negative effect of deer browsing on regeneration in control plots relatively to those without protection.     

Antiproliferative steroidal glycosides from Digitalis ciliata

Two new compounds, a furostanol glycoside (1) and a pregnane glycoside (4), along with eight known compounds, belonging to the classes of spirostane (2,3), pregnane (5–7) and cardenolide (8–10) glycosides, were isolated from the seeds of Digitalis ciliata. Their structures were elucidated by 1D and 2D-NMR experiments as well as ESI-MS analysis. For the first time pregnane glycosides of the diginigenin series have been isolated from D. ciliata. The cytotoxic effects of compounds 1–10 on cell viability of several cancer cell lines, namely human breast cancer (MCF-7), human glioblastoma (T98G), human lung adenocarcinoma (A549), human colon carcinoma (HT-29), and human prostate cancer (PC-3) cell lines were evaluated. Compounds 1, 4, 7 and 8 showed antiproliferative effects against MCF-7, HT-29 and A549 cancer cells with IC₅₀ values ranging from 8.3 to 20 μM. The effects of compounds 1–10 on cell proliferation were evaluated on these three cancer cell lines by cell cycle analysis of DNA content using flow cytometry. Compounds 7, 8 and 10 induced significant changes in G₂/M cell cycle phase of all analyzed cells. The obtained results indicate that compounds 7, 8 and 10 are cytostatic compounds effective in reducing cell proliferation by inducing accumulation of the cells in the G₂/M phase of the cell cycle.     

Does canopy mean nitrogen concentration explain variation in canopy light use efficiency across 14 contrasting forest sites?

The maximum light use efficiency (LUE?=?gross primary production (GPP)/absorbed photosynthetic photon flux density (aPPFD)) of plant canopies has been reported to vary spatially and some of this variation has previously been attributed to plant species differences. The canopy nitrogen concentration [N] can potentially explain some of this spatial variation. However, the current paradigm of the N-effect on photosynthesis is largely based on the relationship between photosynthetic capacity (A(max)) and [N], i.e., the effects of [N] on photosynthesis rates appear under high PPFD. A maximum LUE-[N] relationship, if it existed, would influence photosynthesis in the whole range of PPFD. We estimated maximum LUE for 14 eddy-covariance forest sites, examined its [N] dependency and investigated how the [N]-maximum LUE dependency could be incorporated into a GPP model. In the model, maximum LUE corresponds to LUE under optimal environmental conditions before light saturation takes place (the slope of GPP vs. PPFD under low PPFD). Maximum LUE was higher in deciduous/mixed than in coniferous sites, and correlated significantly with canopy mean [N]. Correlations between maximum LUE and canopy [N] existed regardless of daily PPFD, although we expected the correlation to disappear under low PPFD when LUE was also highest. Despite these correlations, including [N] in the model of GPP only marginally decreased the root mean squared error. Our results suggest that maximum LUE correlates linearly with canopy [N], but that a larger body of data is required before we can include this relationship into a GPP model. Gross primary production will therefore positively correlate with [N] already at low PPFD, and not only at high PPFD as is suggested by the prevailing paradigm of leaf-level A(max)-[N] relationships. This finding has consequences for modelling GPP driven by temporal changes or spatial variation in canopy [N].     

Comparing the intra-annual wood formation of three European species (Fagus sylvatica, Quercus petraea and Pinus sylvestris) as related to leaf phenology and non-structural carbohydrate dynamics

Monitoring cambial phenology and intra-annual growth dynamics is a useful approach for characterizing the tree growth response to climate change. However, there have been few reports concerning intra-annual wood formation in lowland temperate forests with high time resolution, especially for the comparison between deciduous and coniferous species. The main objective of this study was to determine how the timing, duration and rate of radial growth change between species as related to leaf phenology and the dynamics of non-structural carbohydrates (NSC) under the same climatic conditions. We studied two deciduous species, Fagus sylvatica L. and Quercus petraea (Matt.) Liebl., and an evergreen conifer, Pinus sylvestris L. During the 2009 growing season, we weekly monitored (i) the stem radial increment using dendrometers, (ii) the xylem growth using microcoring and (iii) the leaf phenology from direct observations of the tree crowns. The NSC content was also measured in the eight last rings of the stem cores in April, June and August 2009. The leaf phenology, NSC storage and intra-annual growth were clearly different between species, highlighting their contrasting carbon allocation. Beech growth began just after budburst, with a maximal growth rate when the leaves were mature and variations in the NSC content were low. Thus, beech radial growth seemed highly dependent on leaf photosynthesis. For oak, earlywood quickly developed before budburst, which probably led to the starch decrease quantified in the stem from April to June. For pine, growth began before the needles unfolding and the lack of NSC decrease during the growing season suggested that the substrates for radial growth were new assimilates of the needles from the previous year. Only for oak, the pattern determined from the intra-annual growth measured using microcoring differed from the pattern determined from dendrometer data. For all species, the ring width was significantly influenced by growth duration and not by growth rate, which differs from previous studies. The observed between-species difference at the intra-annual scale is key information for anticipating suitability of future species in temperate forests.     

Species proportions by area in mixtures of Scots pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.) and European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.)

Scots pine (Pinus sylvestris L.) and European beech (Fagus sylvatica L.) dominate many of the European forest stands. Also, mixtures of European beech and Scots pine more or less occur over all European countries, but have been scarcely investigated. The area occupied by each species is of high relevance, especially for growth evaluation and comparison of different species in mixed and monospecific stands. Thus, we studied different methods to describe species proportions and their definition as proportion by area. 25 triplets consisting of mixed and monospecific stands were established across Europe ranging from Lithuania to Spain in northern to southern direction and from Bulgaria to Belgium in eastern to western direction. On stand level, the conclusive method for estimating the species proportion as a fraction of the stand area relates the observed density (tree number or basal area) to its potential. This stand-level estimation makes use of the potential from comparable neighboring monospecific stands or from maximum density lines derived from other data, e.g. forest inventories or permanent observations plots. At tree level, the fraction of the stand area occupied by a species can be derived from the proportions of their crown projection area or of their leaf area. The estimates of the potentials obtained from neighboring monospecific stands, especially in older stands, were poorer than those from the maximum density line depending on the Martonne aridity index. Therefore, the stand-level method in combination with the Martonne aridity index for potential densities can be highly recommended. The species’ proportions estimated with this method are best approximated by the proportions of the species’ leaf areas. In forest practice, the most commonly applied method is an ocular estimation of the proportions by crown projection area. Even though the proportions of pine were calculated here by measuring crown projection areas in the field, we found this method to underestimate the proportion by 25% compared to the stand-level approach.