Growth of adult Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica L.) under free-air ozone fumigation

This study attempted to detect the impact of ozone on adult trees of Norway spruce ( Picea abies [L.] Karst.) and European beech ( Fagus sylvatica L.) in an experimental mixed stand in Southern Bavaria, Germany. The aim was to examine whether there is a decrease in growth when trees are exposed to higher than atmospheric concentrations of ozone. This exposure was put into effect using a free-air fumigation system at tree crown level. Growth analysis was carried out on a group of 47 spruce and 36 beech trees, where radial stem increment at breast height - a sensitive index for stress - was measured. The ozone monitoring system allowed values to be obtained for the accumulated ozone exposure (SUM00) of each individual tree, so that their radial increment over three years could be correlated with the corresponding ozone exposure for the same time period. Correlation and regression analysis were then carried out to test the influence of ozone on diameter increment. In both spruce and beech, the initial stem diameter was the most influential factor on radial increment in the following year. A linear model was applied, including the diameter of the preceding year and the ozone exposure of the current year as predicting factors. For spruce trees, a significant negative influence of ozone exposure was found. In contrast, no significant ozone effect on diameter increment of beech was detected. The effect of ozone stress on a large spruce tree can lead to a decrease in potential radial increment of 22 %. The results are discussed in relation to other stress factors such as drought and lack of light.     

Development of biomass proportions in Norway spruce (Picea abies [L.] Karst.)

This study tests the hypotheses that (1) the above-ground structure of Norway spruce (Picea abies [L] Karst.) is derivable from the functional balance theory, and that (2) crown ratio is a key source of structural variation in trees of different age and social position. Twenty-nine trees were measured in three stands (young, middle-aged, and mature), with three thinning treatments (unthinned, normal, and intensive) in the two older stands. There was a strong linear relationship between the total cross-sectional area of branches and that of stem at crown base. Foliage mass was linearly related with stem basal area at crown base. Also an allometric relationship was found between foliage mass and crown length. The mean length (weighted by basal area) of branches obeyed an exponential function of crown length. The parameters of most of these relationships were independent of slenderness (tree height/breast height diameter) and tree age However, total branch cross-sectional area per stem cross-sectional area in the young trees was greater than in the older trees. The young trees also had slightly shorter branches than predicted by the mean branch length equation. This was probably caused by branch senescence which had not yet started in the young stand. The older trees had a relatively long lower crown segment which was growing slowly and senescing. It was proposed that a segmented crown structure is characteristic of shade tolerant tree species, and that the structural model could be further developed by making the two segments explicit.     

Crown development in Norway spruce [<Emphasis Type="Italic"> Picea abies </Emphasis>(L.) Karst.]

This study tests whether crown and stem development in Norway spruce could be described using a modified profile theory. 29 trees from three age-groups (25, 67, 86) with different treatments (unthinned, normally and intensively thinned) were destructively sampled. Crown ratio and crown length varied between age groups and treatments. Crown width was positively correlated with crown length, but branch length along the crown depended on tree age and growing space. Foliage mass density peaked at a relative crown height of 50–70% in middle-aged and mature stands, while young crowns were densest and widest at the base. Foliage mass was predictable from branch and stem cross-sectional area, provided the distance from the top was included. The ratio of foliage mass to branch cross-sectional area increased for 2–4 m down from the tip of the crown, then started to decrease. The relationship between cumulative foliage mass and stem cross-sectional area was non-linear along the stem in the upper crown, but the ratio of cumulative branch to stem cross-sectional area was linear. Trees in the mature and unthinned stands had more cross-sectional area in branches relative to stems than in the young and thinned stands. We conclude that the profile theory needs modification regarding (1) crown shape which varies with age and growing space, and (2) the ratio of foliage mass to branch area which varies along the stem. Both aspects emphasise the need to include impacts of disuse of sapwood pipes in models of crown and stem development.     

Relationships between tree dimension and coarse root biomass in mixed stands of European beech (Fagus sylvatica L.) and Norway spruce (Picea abies[L.] Karst.)

Relationships between tree parameters above ground and the biomass of the coarse root system were examined in six mixed spruce-beech stands in the Solling Mountain region in northwest Germany. The selected stands were located on comparable sites and covered an age range of 44 to 114 years. Coarse roots (d?\ge?2 mm) of 42 spruce and 27 beech trees were sampled by excavating the entire root system. A linear model with logarithmic transformation of the variables was developed to describe the relationship between the coarse root biomass (CRB, dry weight) and the corresponding tree diameter at breast height (DBH). The coefficients of determination (R ²) attained values between 0.92 for spruce and 0.94 for beech; the logarithmic standard deviation values were between 0.29 and 0.43. A significantly different effect of tree species on the model estimates could not be detected by an analysis of co-variance (ANCOVA). For spruce, the derived relationships were similar to those reported in previous studies, but not for beech. Biomass partitioning in the tree compartments above and below ground differs significantly between spruce (coarse root/shoot ratio 0.16±0.06) and beech (coarse root/shoot ratio 0.10±0.03) in the mixed stands. These results are similar to those given in other studies involving pure spruce and beech stands on comparable sites in the region, although the ratios of pure stands in other regions growing under different site conditions are somewhat higher. Comparing trees of the same DBH classes, root/shoot ratios of spruce are 1.2 to 3 times higher than those of beech. Dominant spruce trees (DBH>60 cm) attained the highest ratios, suppressed beech trees (DBH<10 cm) the lowest. Site conditions of varying climate and soils and interspecific tree competition are likely to affect root/shoot ratio and DBH-coarse root biomass relationships. The greater variability in beech compared with spruce indicates a high 'plasticity' and adaptability of beech carbon allocation. Thus, the derived equations are useful for biomass estimates of coarse roots involving trees of different ages in mixed stands of spruce and beech in the Solling Mountains. However, application of these relationships to stands in other regions would need further testing.     

Structural crown properties of Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica [L.]) in mixed versus pure stands revealed by terrestrial laser scanning

How tree morphology develops in mixed-species stands is essential for understanding and modelling mixed-stand dynamics. However, research so far focused on the morphological variation between tree species and neglected the variation within a species depending on intra- and interspecific competition. Our study, in contrast, addresses crown properties of nine mature Norway spruces (Picea abies [L.] Karst.) of a pure stand and compares them with ten spruces growing in mixture with European beech (Fagus sylvatica [L.]). The same was done with 11 pure stand beeches and 12 beeches growing in mixture with spruce. Through application of a terrestrial laser scanner and a new skeletonization approach, we deal with both species’-specific morphological traits such as branch angle, branch length, branch bending, crown volume and space occupation of branches within the crown, some of which were hardly accessible so far. Special attention is paid to distinct differences between trees growing in mixed and pure stands: for spruce, our study reveals significantly longer branches and greater crown volumes in the mixed stand when compared to the pure stand. In case of European beech, individuals growing in mixture show flatter branch angles, more distinct ramification, greater crown volumes and a lower share of a single branch’s space occupation in the total crown volume. The results show that the presented methods yield detailed information on the morphological traits analyzed in this study and that interspecific competition on its own may have a significant impact on crown structures. Implications for production ecology and stand dynamics of mixed-species forests are discussed.     

Climate effects on productivity and resource-use efficiency of Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica [L.]) in stands with different spatial mixing patterns

Species mixing is widely held to stabilize productivity, increase resilience and contribute to risk minimization in forest stands in need of special as a result of longevity. However, research on the effects of mixing on productivity and resource consumption so far yielded fairly incoherent results rather than general findings. We focused on the effects of the spatial mixing pattern and the annual climate conditions on the mixing effect, which to date have seldom been considered as modifiers of mixing effects. Nine years of intensive survey of four pure stands of Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica [L.]) and two mixed plots with different mixing pattern showed: (1) mixing hardly changed annual net primary productivity at stand level when Norway spruce and European beech are cultivated group-wise but increased by 37 % on account of a higher efficiency of water and light use in individual tree-wise mixture. (2) Favourable climatic conditions increased the superiority of mixed versus pure stands productivity, while, in particular, water stress cancelled the benefit of mixing considerably. (3) An interaction between the spatial pattern and variable climatic conditions was revealed. Both improved light and water use were found in favourable years in close inter-specific intermingling. However, in unfavourable years the spatial pattern played a less pronounced role in terms of productivity.     

Mixed Norway spruce (Picea abies [L.] Karst) and European beech (Fagus sylvatica [L.]) stands under drought: from reaction pattern to mechanism

Key message

We review causes of synergies in mixed-species stands, develop guiding hypotheses for revealing their mechanisms and present a rainfall exclusion experiment along with a transect (KROOF) for exploring drought effects.

Abstract

While monocultures have dominated forest research and practice in the past, in face of growing resource scarcity and climate change, mixed-species stands are on the advance. Long-term observations show that mixed-species stands frequently over-yield monocultures, and they further suggest that the over-yielding is often higher on poor than on fertile sites and in low-growth than in high-growth years. However, the underlying causes have not yet been clarified. We start with a review of, among others, hydraulic redistribution, complementary eco-physiological traits, and ectomycorrhizal networks as possible causes behind the observed productivity gains in mixed-species stands. Then, we develop guiding hypotheses for further exploration of synergies in mixed-species stands. Finally, we introduce into the concept and model approach of the rainfall exclusion experiment for examining the role of water supply in mixed versus pure forest stands of spruce and beech. At the Kranzberg site, six plots are equipped with water retention roofs, which only close during rain events. The remaining six plots serve as non-roofed control. Together with the rainfall exclusion experiment, an ecological gradient with five sites extending through SE-Germany forms the “Kranzberg Roof Experiment” (in short KROOF). Kranzberg Forest is a part of this gradient from moist to dry conditions, with each site providing mixed and pure associations each of spruce and beech. The ecophysiological approach will be complemented by tree ring analysis and modelling of productivity of the tree associations under intense drought.     

Structure and stand development in three subalpine Norway spruce (Picea abies (L.) Karst.) stands in Paneveggio (Trento, Italy)

1. Three permanent plots (100×0 m) were established in the subalpine Norway spruce (Picea abies (L.) Karst.) forest of Paneveggio in the spring of 1993, to begin a long-term forest ecosystem research project. The main purpose of these plots was to provide information about subalpine Norway spruce stand dynamics and to provide suggestions for close-to-nature silviculture. 2. The three stands were selected to represent the most common forest structures in the Paneveggio forest. The first stand is close to forestry roads, has a relatively regular and continuous canopy, and thinning and cutting operations only ended in the 1980s; the second stand is far from forest roads and has developed without anthropogenic influence for several decades; the third one is located at the present upper limit of the pure spruce forest and, apparently, was heavily used in the past as a pasture. 3. The first step in the investigation was to describe the structure and to study the history of the three stands using both written evidence from manage- ment plans and biological archives from tree rings. 4. The stands in plots 1 and 2 began to establish after a disturbance that removed part of the previous stands according to dendroecological studies, which are partially supported by written evidence. The remaining parts of these stands were eliminated by two major disturbances that occurred during the following decades. Written records about the use of the forest lead us to assume that the initial disturbances that occurred in the two stands were logging activities as a part of a group shelterwood system. The stand in plot 2 has developed without significant human interference for about half a century as confirmed by the presence of many dead trees. The stand in plot 3 consists of old trees that were part of an open stand and a secondary population that established after cessation of grazing. 5. The study has confirmed that dendroecological techniques can be used to identify occurrence and intensity of previous disturbance in forests stands, although at Paneveggio it is difficult to distinguish between natural and anthropogenic disturbances in the tree ring record. The presence of human activity necessitates investigation of multiple lines of evidence. 6. Paneveggio's forest management plans were useful in the interpretation of the data obtained through dendroecological analysis, although events did not always correspond because data from the management plans (yearly thinning, felling, wind-throw damage) never gave stand-level details, but applied to areas of several hectares. Despite these limitations, the information included in the management plans is of crucial importance in studying stand history and only by using all these sources of information is it possible to delineate the most important features of the history and disturbance that affected the origin and subsequent growth of the forest stands.     

In-situ soil solution chemistry in an acid forest soil as influenced by growing roots of Norway spruce (Picea abies [L.] Karst.)

A new approach for non-destructive monitoring of soil solution chemistry in high spatial and temporal resolution for rhizosphere studies is presented. In a 5×10 mm grid, 30 micro suction cups (1mm) were installed in a rhizotron with Norway spruce (Picea abies [L.] Karst.) growing in low pH B-horizon soil. Roots grew through the grid, closely passing the suction cups. Soil solution composition before, during and after root passage was determined. For K⁺ and Mg²⁺ a significant decrease of soil solution concentration near root tips and elongation zones was observed, indicating a marked uptake of these elements. Mg²⁺ concentration was also significantly lowered when the root system aged, suggesting that this ion might also be taken up in older parts of the root system. No influence of growing roots was found on Na⁺-concentrations.     

Seasonal development of the photosynthetic performance of Norway spruce (Picea abies [L.] Karst.) under magnesium deficiency

In order to investigate the influence of different magnesium nutrition on photosynthesis, one hundred 6-year-old spruce trees derived from one clone were planted in October 1990 into a special out-door experimental construction, where they were cultivated in sand culture with an optimal supply of nutrients, except magnesium, via circulating nutrient solutions. Magnesium was added to the nutrient solutions in three different concentrations, varying from optimal to severe deficient supplies. During the first vegetative period in 1991, photosynthetic performance and carboxylation efficiency were measured under saturating light, controlled CO₂ conditions, optimal temperature and humidity, using a minicuvette system.During summer, the trees under moderate magnesium deficiency developed tip yellowing symptoms on older needles, while the youngest needles remained green with unchanged chlorophyll contents. Trees under severe magnesium deficiency showed yellowing symptoms on all needle age classes combined with decreased chlorophyll contents in the youngest needles as well. In comparison with the controls, the photosynthetic performance of the 1-year-old needles was significantly lower in both deficiency treatments. The same was observed in the youngest needles of the trees under severe deficiency. Trees under moderate deficiency treatment decreased in photosynthetic performance during the summer without reduction of chlorophyll contents. The reduction of photosynthetic rates corresponded to a decrease in carboxylation efficiency, which is taken as a measure of the activity of the enzyme ribulose-1,5-bisphosphate carboxylase. This reduction, together with the observed increase of carbohydrate contents in needles of trees growing under magnesium deficiency, led to the assumption that the photosynthetic carbonfixation is reduced as a consequence of the accumulation of carbohydrates.     

Influence of UV-B radiation on photosynthetic activity and chlorophyll fluorescence kinetics in Norway spruce [Picea abies (L.) Karst.] seedlings

Norway spruce [Picea abies (L.) Karst.] seedlings were grown in greenhouses with two supplemental levels of ultraviolet-B (UV-B) radiation. Photochemical efficiency of photosystem II and vitality index were determined monthly. At the end of the experiment, growth, chlorophyll content and photosynthetic rates were measured. The data indicate that low temperature in winter affected light dependent processes in experimental plants including control, while the rise of ambient temperatures, moderate this effect. The synergistic effects of UV-B radiation and low temperatures could only be observed in the second winter period. Measurements of net photosynthetic activity in the second winter period showed significant differences between treated and untreated plants.     

Seasonal accumulation of ultraviolet-B screening pigments in needles of Norway spruce (Picea abies (L.) Karst.)

Conifer needles are highly effective in screening ultraviolet-B radiation (280–320 nm). This ability is mainly attributed to the presence of flavonoids and hydroxycinnamic acids in the epidermal tissue. In two field cabinet experiments with two different clones of Norway spruce we assessed the seasonal accumulation of UV-B screening pigments under near-ambient, and close-to-zero UV-B irradiation. At the beginning of needle development, i.e. in June, kaempferol 3- O -glucoside was the dominant UV-B screening pigment. It was replaced during needle differentiation by the more effective diacylated flavonol glucosides, particulary kaempferol 3- O -(3",6"- O -di- p -coumaroyl)-glucoside, which reached highest concentrations in July. In addition to the soluble pool of diacylated flavonol glucoside derivatives, a cell wall-bound UV-B screen in the epidermal cell walls was formed during needle differentiation, consisting mainly of p -coumaric acid and kaempferol 3- O -glucoside. An effect of UV-B radiation on the accumulation of diacylated flavonol glucosides was only observed in 1996 with clone 2, when the concentrations of kaempferol 3- O -(3",6"- O -di- p -coumaroyl)-glucoside were significantly higher in July and August under field, and near-ambient than under close-to-zero UV-B irradiance. For wall-bound p -coumaric acid and kaempferol 3- O -glucoside UV-B radiation enhanced the concentrations of these compounds by approximately 20% in relation to the concentrations in close-to-zero UV-B-treated plants in both field cabinet experiments.     

Time change of aluminium toxicity in the acid bulk soil and the rhizosphere in Norway spruce (Picea abies (L.) Karst.) and beech (Fagus sylvatica L.) stands

Context

In acidic forest soils, aluminium can alter tree health due to its potential toxicity. Aluminium phytotoxicity is mainly influenced by its chemical form and its availability.

Methods

As physical-chemical indicators of Al toxicity in soil, Al speciation in soil solutions and in the exchange complex was measured in the rhizosphere and the bulk soil of two tree species (Norway spruce (Picea abies (L.) Karst.) and European Beech (Fagus sylvatica L.) in an acidic soil and in 4 months (November, February, May and August) representing the four seasons in a year.

Results

In the bulk soil, Al toxicity was generally higher under Norway spruce than under beech. Furthermore, temporal changes in Al behaviour were identified under Norway spruce but not under beech. The monomeric Al in the soil solutions and the exchangeable Al in the solid soil increased significantly in February under Norway spruce and were positively correlated with nitrate concentration, suggesting that nitrate influence Al speciation and mobility under Norway spruce. In the rhizosphere, Al toxicity was restricted through Al complexation by organic compounds and by nutrient contents independently from the season. The ecological importance of the rhizosphere in Al detoxification is discussed.

Conclusions

This study suggests that plant specific differences as well as seasonal changes in plant physiology, microbial activity and microclimatology influence aluminum toxicity in acid forest soils.     

The dependency of the size-growth relationship of Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica [L.]) in forest stands on long-term site conditions, drought events, and ozone stress

Against a backdrop of increasing climate change, the effects of site conditions, drought events and ozone stress on the size-growth relationship in Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica [L.]) stands are analyzed. The size-growth relationship is represented by a straight line defined by intercept and slope of a simple linear equation with stem diameter at height 1.30 m as independent variable and annual stem diameter increment at height 1.30 as dependent variable. On the basis of 64 long-term experimental plots dating back to 1871 and representing an ecological gradient from fertile to poor sites, it is shown that poorer sites exhibit shallower slopes of the linear size-growth relationships than fertile sites. Annual measurements of the size-growth relationship, including the extremely dry years of 1976 and 2003, also showed that lower stand growth rates result in shallower size-growth relationship slopes. By comparing stands with and without experimental twice-ambient ozone exposure between 2000 and 2007, it was found that ozone stress can significantly reduce the slope of the size-growth relationship. This indicates that limiting site condition, whether acute or chronic in nature, distinctly reduces the superiority of tall trees, and that a lower degree of resource limitation increases the steepness of the size-growth relationship. The causes for this behavior and the consequences for stand dynamics, silvicultural treatment and prognostication by models are discussed.     

Influence of cultivation method on root grafting in Norway spruce (Picea abies (L.) Karst.)

Root grafting is the process by which a functional union of two or more roots subsequent to their formation is formed. The above- and below-ground parts of three Norway spruce stands (natural stand, Umbric Luvisol; row-culture and group-culture, Planosol; stand ages 40, 43 and 43, respectively) of high site quality (I) were investigated. Stand densities were 1550, 1783 and 1722 stems ha^-1, respectively. In all investigated stands, root grafting was most sensitive to tree spacing. Grafts were observed in case the distance between the trees was 0.7–1.2 m. Grafts occurred always in areas of higher rooting density, in a row of the row-culture and within a tree group in the group-culture. Root grafting was enhanced in case of a narrower humus horizon in the group culture compared with the row-culture, 16.5 and 30 cm, respectively; the humus horizon contained 99% and 95% of conducting roots with d ≥ 5 mm, respectively. Root graftings occurred in 75% of excavated trees in the group-culture, in 37.5% of excavated trees in the row-culture and in 33.3% of excavated trees in the natural stand. Stand age was 24 years in the row-culture and 22 years in the group-culture at the beginning of root grafting. No grafts occurred between two suppressed trees, whereas in 86–100% of all cases, at least one tree was dominant or codominant. In row- and group-cultivated Norway spruce stands, the initial minimum diameter of the grafted root without bark was from 1 to 3 cm in 63% of cases. Grafting of roots with d < 1 cm or d > 10 cm was rare or absent. Root grafting had usually begun at the root age of 10–20 years (46% of cases). This revised version was published online in June 2006 with corrections to the Cover Date.