Amino nitrogen and phenolic constituents of bark of American beech,Fagus grandifolia,and infestation by beech scale,Cryptococcus fagisuga

Concentrations of amino acids, total amino nitrogen, and phenols, ratio of phenol to amino nitrogen, and pH in the bark of American beech, Fagus grandifolia, were determined to see if these chemical constituents were correlated with susceptibility of trees to infestation by the beech scale, Cryptococcus fagisuga. The relationship of these bark constituents with scale infestation levels (none, light, and moderate), tree size (large vs. small trees), and bark shading was determined. Levels of some individual amino acids and total amino acid content in moderately infested trees were significantly higher than in uninfested trees. Large uninfested beech trees, which tend to be more readily infested by scale, had significantly higher concentrations of aspartic acid. Concentrations of phenol in the outer bark of both infested and uninfested trees were significantly higher than in the inner bark. However in bark infected by Nectria, phenol levels in the inner bark were significantly higher than in the outer bark.     

Bark phenolics of American beech (Fagus grandifolia) in relation to the beech bark disease1

The amount of total extractable phenols was determined for bark sections obtained from behind cankers naturally induced by Nectria coccinea var. faginata and from behind mechanically inflicted wounds on stems of American beech. Healthy bark from susceptible trees was found to contain a similar level of phenolics as bark from trees determined to be resistant to the beech bark disease. Six months after wounding, wound-altered bark from susceptible trees was found to be lower in phenols than woundaltered bark from resistant trees. Inoculation of wounds with N. coccinea var. faginata resulted in decreased phenolic levels in bark sections nearest the wound surface, and increased phenolic levels in sections nearest the vascular cambium, several mm distant. Phenolic levels in injured or infected bark appear to follow similar patterns as those resulting from injury or infection of xylem tissues.     

Scale insects,decay and canker fungi in American beech

Tripartite interactions among phytophagous insects, pathogens and their host plants provide insight into the role of host physiology in determining susceptibility to attack. American beech (Fagus grandifolia) often is simultaneously attacked by beech scale (Cryptococcus fagisuga), one or more Neonectria pathogens and Xylococculus betulae that can result in beech bark disease (BBD). Additionally, beech is frequently infected by heartrot‐decay fungi. Cursory observations in 2011 suggested that beech scale and Neonectria lesion densities were lower and greater, respectively, on trees with decay. In 2012, digital image analysis was used to quantify densities of these organisms on 123 beech from the Adirondack region of New York. Three groups of study trees (n = 41) were used: Inonotus glomeratus‐infected, Phellinus igniarius‐infected and non‐decay trees. Trees infected by decay pathogens supported lower densities of beech scale and higher densities of Neonectria. Densities of X. betulae did not significantly vary among decay groups. These results may be explained by decay‐induced changes in host physiology. Additional work is needed to elucidate the potential role of host bark chemistry in the BBD complex.     

Gap dynamics in the Western Carpathian mixed beech old-growth forests affected by spruce bark beetle outbreak

Understanding how species-specific disturbances affect the dynamics of mixed forests is becoming increasingly important due to rapidly changing disturbance regimes. This study estimated the effect of Norway spruce (Picea abies (L.) Karst.) mortality on the disturbance processes in two mixed beech stands of the Western Carpathians that were affected by a bark beetle outbreak. We evaluated the size distribution, fraction of canopy and expanded gaps, the characteristics of gapmakers and the contribution of different species to gap size. The measured canopy gap fraction was <5%, and most canopy gaps were small (<100 m²). Spruce was the most abundant gapmaker, and its share among gapmakers was 3–6 times higher than its share in the canopy. We found that the increase in spruce mortality due to the outbreak resulted in a fine-scale mortality pattern. However, spruce gapmakers did not contribute much to gap area size, as shown by a weak correlation between the number of spruce gapmakers and the area of expanded gaps. Diameter distribution of living versus recently dead trees showed that beech mortality occurred disproportionately in large size classes. However, dead spruce trees were equally frequent in all diameter classes, which means beetles did not exclusively attack larger trees in these stands during the outbreak. We conclude that spruce mortality may have influenced successional processes by giving a competitive advantage to two other species that were not affected by the outbreak, provided that a high deer browsing intensity does not hinder the regeneration of new seedlings.

     

Relationships among beech bark disease,climate, radial growth response and mortality of American beech in northern Maine,USA

A majority of beech forests across Maine first experienced beech bark disease (BBD) from 1935 to 1960 when sap feeding by an introduced beech scale insect, Cryptococcus fagisuga, allowed lethal fungal infections primarily by Neonectria ditissima and/or Neonectria faginata. Beech stands along the Maine–Quebec border in northern Maine were excluded from this initial killing phase presumably due to cold winter temperatures that inhibited scale survival. However, a sharp increase in beech mortality after 2002 occurred in previously uninfected border stands and stands long affected by BBD. Beech mortality averaged 50% across northern Maine during 2003–2006. To identify plausible stresses that could explain the mortality, a dendropathological study was conducted from 2005 to 2006 in northern Maine that quantified temporal and spatial relationships between possible stressors with beech mortality and growth decline. Nineteen sets of high‐ and low‐mortality plots were located randomly across four bioregions. Increment cores were taken from both beech trees (n = 565) and associated tree species (n = 450). A growth change index of increments was used to evaluate beech responses to biotic and climatic stresses. A prolonged period of relatively mild winters without temperatures lethal to scale insect (Neonectria was found infecting weakened trees across the region. Drought, beech scale and Neonectria are plausible explanations for the episode of high beech mortality in northern Maine. This is the first report of a major killing phase of beech within the BBD ‘aftermath’ forests.     

Fate of nitrogen released from 15N-labeled litter in European beech forests

The decomposition and fate of 15N-labeled beech litter was monitored in three European beech (Fagus sylvatica L.) forests (Aubure, France; Ebrach, Germany; and Collelongo, Italy) for 3 years. Circular plots around single beech trees were isolated from roots of neighboring trees by soil trenching, and annual litterfall was replaced by 15N-labeled litter. Nitrogen was continuously released from the decomposing litter. However, over a 2-year period, this release was balanced by the incorporation of exogenous N. Released N accumulated mainly at the soil surface and in the topsoil. Microbial biomass remained almost constant during the experiment at all sites except for considerably lower values at Ebrach. The 15N enrichment of the microbial biomass increased strongly during the first year and then remained stable. The 15N released from the decomposing litter was rapidly detected in roots and leaves of the beech trees, increasing regularly and linearly over the course of the experiment. The uptake of litter-released 15N by the trees was reduced under conditions that reduced tree growth. Under these conditions, leaves and fine roots were the dominant N sinks, and little N was allocated to other plant parts. By contrast, N uptake and N allocation from leaves to stem and bark tissues increased when tree growth was enhanced. Budgets for 15N showed that 2 to 4% of litter-released N was incorporated into the trees, about 35% remained in the litter and about 50% reached the topsoil.     

Some Aspects of the Growth of Beech Trees and the Incidence of Beech Bark Disease on Chalk Soils

Growth characteristics and the incidence of beech bark diseasewere assessed within beech stands, aerial photographs of whichshowed discrete areas of foliar chlorosis. Chlorosis was mainlyconfined to distinct soil zones which contained microscopicallydivided chalk. Compared with the non-chlorotic trees, thosetrees growing in the chalky soil type had smaller mean diameterand had a higher current stocking density. The incidence ofthe early (insect infestation) phase of beech bark disease wasthe same on the two soil types, but the second phase (bark necrosis)was more severe on the chalky soil.     

Modelling the economically viable wood in the crown of European beech trees

Long-term forest development programs in Germany aim on an increase of close-to-nature broadleaf forest stands. This means that the economic importance of European beech is expected to increase. The economic potential of a tree basically consists of the stem as well as the economically viable wood volume in the crown. Due to the high morphological variability of European beech crowns, taper models are often not satisfactory for predicting the economically viable wood volume arising from crowns. Prediction models with a higher precision are recently still lacking. Aim of this study is thus the development of prediction model for the economically viable crown wood volume of European beech trees.We determined the distribution of the wood volume in the crown over the branch diameters using the multistage ‘randomized branch sampling’ method (RBS). The tree-specific wood volume distribution on the branch diameters were used to cluster all sampled trees into 3 groups. Additionally, we developed a method able to distinguish between economically viable and unviable crown branches. Basing on the RBS measurements as well as revenues and processing costs, we modeled the economically viable wood volume from the crown for each tree. To calculate the wood volume under bark, we parameterized a bark thickness function from disk samples of the trees.We showed that the European beech crowns could be clustered into 3 groups differing in their wood volume distribution. The economically viable wood volume in the crown significantly depended on this grouping parameter as well as diameter at breast height (DBH). By contrast, the total amount of wood in the crown only depended on DBH. The differing viable wood volumes in the crowns were thus explained by different wood distributions and not by differing total crown wood volume. To make the results applicable in practice forestry, the modelling results were used to develop a regression formula able to predict the economically viable wood volume in the crown depending on the DBH and the crown type. As the crown type can also be predicted via measurable tree covariates, the regression model of the viable wood volume in the crown can be used as a support tool for the management of European beech stands. Sensitivity analysis quantifies how harvest revenues and costs translate into different viable tree volume.     

Canopy recruitment in the beech (<Emphasis Type="Italic">Fagus engleriana</Emphasis>) forest of Mt. Shennongjia,Central China

A dendrochronological technique was used to investigate canopy recruitment of beech (Fagus engleriana Seem.) trees in western Hubei Province of Central China by identifying growth releases through gap formation. The results indicated that 83% and 17% of the canopy trees regenerate in gaps and the understory, respectively. Radial growth analyses demonstrated that there are three modes for beech regeneration and canopy recruitment: (1) gap origin without suppression; (2) gap origin with period(s) of suppression; and (3) understory origin. Multiple periods of release and suppression imply that beech regeneration and canopy recruitment are very sensitive to gap formation.     

Radial distribution of carbohydrate reserves in the trunk of declining European beech trees (Fagus sylvatica L.)

Context

European beech (Fagus sylvatica L.) is considered threatened by anticipated climate change, but the physiological causes of potential beech decline or mortality remains poorly understood.

Aims

The purpose of the present study was to fuel debate about the assumption that carbohydrate depletion is involved in the decline of mature European beech.

Methods

The health status of beech trees from a severely declining stand was visually assessed by examining their crown condition. Content and radial distribution of non-structural carbohydrates (starch and soluble carbohydrate) were analyzed in the trunks and compared to those reported earlier in trunks of healthy beech trees.

Results and discussion

The distribution of carbohydrate in the beech trunks recorded here seemed affected by decline. We found a stronger radial decrease of starch content than those reported earlier for healthy beech trees. Carbohydrate reserves appear partially maintained in the outermost rings while starch depletion occurred in older wood rings in declining trees that may be able to mobilize carbohydrate reserves from older wood rings in response to successive climatic constraints.     

Micromorphology of naturally degraded beech and spruce barks

The micromorphological changes occurring in the bark tissues of beech and spruce trees during natural degradation in the forest have been investigated and compared with those obtained previously using wood-decaying fungi. The relevance of the bark degradation process for the environment is also briefly discussed.     

Influence of nitrogen fertilization on minerals, carbohydrates, amino acids and phenolic compounds in beech (Fagus sylvatica L.) leaves

Beginning in 1985, plots in a 120-year-old beech (Fagus sylvatica L.) stand in the province of Scania, southern Sweden were fertilized annually with ammonium nitrate. In August 1989, leaves from both fertilized and unfertilized trees were analyzed for mineral nutrients, soluble carbohydrates, amino acids and phenolic compounds. Leaf concentrations of total nitrogen were increased by N fertilization. Ammonium was not detected in the leaves, but NO(3) (-) was increased by up to 65% as a result of N fertilization. Nitrogen fertilization decreased foliar concentrations of P and Cu, leading to a nutrient imbalance relative to N. Of the free amino acids, glutamic acid and aspartic acid predominated and, together with glutamine and asparagine, were significantly increased by N fertilization. The concentrations of amides were 2-4 times higher in fertilized trees than in unfertilized trees. Soluble carbohydrates did not change in response to N fertilization, but total phenolic compounds decreased markedly. The reduced concentration of phenolics may explain the greater susceptibility to parasite attacks of trees exposed to N deposition and excess N.     

Relationship between branch-scar parameters and knot features of oriental beech (Fagus orientalis Libsky)

The classification of roundwood is inextricably linked to the measurement of a particular single wood defect. The appearance, location, and number of defects are important in the quality evaluation of logs and sawn timber, and the most important defects are knots. The purpose of this study was to investigate the relationship between the appearance of branch scars and features of the related knot inside oriental beech logs, and to model the relationship between well-defined branch-scar and knot parameters. One hundred and fifty knots in 15 stems of oriental beech trees were studied. Image analysis software was used to measure the branch-scar and knot features. The results showed a significant positive correlation between the branch-scar parameter “moustache length” and the knot length. The ratio of branch-seal length to width was found to be a good estimator of the stem diameter at the time of knot occlusion and the amount of clear wood between the knot occlusion and the bark. The relationship obtained for the oriental beech stem radius at time of knot occlusion confirms relationship reported for European beech (Fagus sylvatica L.).     

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.     

Some factors affecting canker formation in American beech inoculated with Nectria coccinea var. faginata1

Combinations of Nectria coccinea var. farinata (the primary causal agent of beech bark disease), Fusarium oxysporum, and associated bacteria were inoculated into American beech trees at two sites in New Hampshire. Neither F. oxysporum nor bacteria appeared to affect the size of cankers formed. Canker size was significantly related to bark thickness at one site when only one mating type of the pathogen was inoculated alone.     

Beech decline in Central Europe driven by the interaction between Phytophthora infections and climatic extremes

During the past decade, and in particular after the wet year 2002 and the dry year 2003, an increasing number of trees and stands of European beech (Fagus sylvatica L.) in Bavaria were showing symptoms typical for Phytophthora diseases: increased transparency and crown dieback, small‐sized and often yellowish foliage, root and collar rot and aerial bleeding cankers up to stem heights of >20 m. Between 2003 and 2007 134 mature beech stands on a broad range of geological substrates were surveyed, and collar rot and aerial bleeding cankers were found in 116 (86.6%) stands. In most stands the majority of beech trees were declining and scattered or clustered mortality occurred. Bark and soil samples were taken from 314 trees in 112 stands, and 11 Phytophthora species were recovered from 253 trees (80.6%) in 104 stands (92.9%). The most frequent species were P. citricola, P. cambivora and P. cactorum. Primary Phytophthora lesions were soon infected by a series of secondary bark pathogens, including Nectria coccinea, and wood decay fungi. In addition, infected trees were often attacked by several bark and wood boring insects leading to rapid mortality. Bark necroses were examined for their probable age in order to determine whether the onset of the current Phytophthora epidemic was correlated to rainfall rates recorded at 22 Bavarian forest ecosystem monitoring stations. A small‐scale survey in nine Bavarian nurseries demonstrated regular infestations of all beech fields with the same range of Phytophthora species. The results indicate that (1) Phytophthora species are regularly associated with beech decline and may also be involved in the complex of ‘Beech Bark Disease’, (2) excessive rainfalls and droughts are triggering the disease, and (3) widespread Phytophthora infestations of nursery stock might endanger current and future silvicultural projects aiming on the replacement of non‐natural conifer stands by beech dominated mixed stands.     

Soluble N compound profiles and concentrations in European beech (Fagus sylvatica L.) are influenced by local climate and thinning

We assessed seasonal changes of total soluble nonprotein nitrogen compounds (TSNN) in adult European beech trees (Fagus sylvatica, L.) growing under different local climate during the growing season immediately following a thinning treatment and 3 years later. In both years, samples of leaves, xylem sap and phloem exudates from beech trees growing in thinned and unthinned (control) stands on a dry, warm SW exposed and a cooler, moist NE exposed site were collected in May, July and September. In May of both years, asparagine (Asn) and glutamine (Gln) were most abundant in leaves and xylem, respectively, whereas arginine (Arg) dominated in the phloem. In July, TSNN concentrations decreased in all tissues and sites, but differences in water availability between aspects were reflected in TSNN concentrations. In September, differences in the increase of Arg concentration in the phloem were related to differences in the onset of senescence between treatments. Thinning treatment increased amino compound concentrations of beech tissues in July on both aspects, particularly at the NE thinned site. It is supposed that, the N balance of adult beech is favoured by both, the thinning treatments as well as the cool-moist climate prevailing at the NE aspect.     

Physiological defence reactions of young beech trees (Fagus sylvatica) to attack by Phyllaphis fagi

The physiological reaction to attack by the lachnid Phyllaphis fagi compared with non-infested control plants was examined in whole leaves and leaf phloem of beech (Fagus sylvatica) saplings as well as in beach seedlings cultivated in different nutrient substrates. The colonisation density of the lachnids on beech seedlings increased with improved nutrient supply. However, growth was not depressed to a comparable extent. Defence reactions were found at three hierarchic levels. Infested plants exhibited a strongly decreased amino acid/monosaccharide ratio in the entire leaf and leaf phloem. In addition, secondary plant defence substances were detected in beech saplings and lamma shoot leaves of beech seedlings. A systematic reaction, dependent on the infestation density of the lachnid was observed in the leaf phloem of the beech saplings.     

Competition modifies effects of enhanced ozone/carbon dioxide concentrations on carbohydrate and biomass accumulation in juvenile Norway spruce and European beech

Elevated concentrations of carbon dioxide ([CO2]) and ozone ([O3]) affect primary metabolism of trees in opposite ways. We studied their potential interactions on carbohydrate concentrations and contents. Two hypotheses currently under debate were tested. (1) Stimulation of primary metabolism by prolonged exposure to elevated [CO2] does not compensate for the adverse effects of O3 on carbohydrate accumulation and biomass partitioning to the root. (2) Growth in a mixed-species planting will repress plant responses to elevated [O3] and [CO2] relative to conditions in a monoculture. To this end, European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst.) saplings grown under conditions of intra- and interspecific competition were pre-acclimated for 1 year to ambient or elevated [CO2]. In the following 2-year phytotron study, trees were exposed to factorial combinations of ambient and elevated [O3] and [CO2]. The total carbohydrate content (sugar and starch) of spruce was greater in plants exposed to elevated [CO2] than in plants exposed to ambient [CO2]. In beech, the opposite response was observed, especially when this species was grown in combination with spruce. Overall, the data did not support Hypothesis 1, because the adverse effects of O3 were counteracted by elevated [CO2]. Support for Hypothesis 2 was species-dependent. In beech saplings, reduction of carbohydrates by elevated [O3] and stimulation by elevated [CO2] were repressed by competitive interaction with spruce. In contrast, in spruce, stimulation of carbohydrates by elevated [CO2] was similar in mono- and mixed cultures. Thus Hypothesis 2 was supported for beech but not spruce. We conclude that, in juvenile beech and spruce, a 3-year exposure to elevated [CO2] counteracts the adverse effects of O3 on carbohydrate concentrations and contents. For beech, sensitivity to elevated [CO2] and [O3] was high in monoculture but was largely repressed by interspecific competition with spruce. In contrast, the response of spruce to perturbations of atmospheric chemistry was not significantly affected by either intra- or interspecific competition.     

Contrasting distribution and seasonal dynamics of carbohydrate reserves in stem wood of adult ring-porous sessile oak and diffuse-porous beech trees

We tested the hypothesis that broad-leaved forest species with contrasting wood anatomy and hydraulic system (ring-porous versus diffuse-porous) also differ in distribution and seasonal dynamics of carbohydrate reserves in stem wood. Total nonstructural carbohydrate (TNC) reserves (starch and sugars) were measured enzymatically in the 10 youngest stem xylem rings of adult oak (Quercus petraea (Matt.) Liebl.) and beech (Fagus sylvatica L.) trees during an annual cycle. Radial distribution of carbohydrates was investigated according to ring age. On all dates, oak trees had twofold higher TNC concentration than beech trees (41 versus 23 mg g(DM)(-1)), with starch accounting for the high TNC concentration in oak. Seasonal dynamics of TNC concentration were significantly (P < 0.05) more pronounced in oak (20-64 mg TNC g(DM)(-1)) than in beech (17-34 mg TNC g(DM)(-1)). A marked decrease in TNC concentration was observed in oak trees during bud burst and early wood growth, whereas seasonal fluctuations in TNC concentrations in beech trees were small. The radial distribution of TNC based on ring age differed between species: TNC was restricted to the sapwood rings in oak, whereas in beech, it was distributed throughout the wood from the outermost sapwood ring to the pith. Although the high TNC concentrations in the outermost rings accounted for most of the observed seasonal pattern, all of the 10 youngest xylem rings analyzed participated in the seasonal dynamics of TNC in beech trees. The innermost sapwood rings of oak trees had low TNC concentrations. Stem growth and accumulation of carbon reserves occurred concomitantly during the first part of the season, when there was no soil water deficit. When soil water content was depleted, stem growth ceased in both species, whereas TNC accumulation was negligibly affected and continued until leaf fall. The contrasting dynamics and distribution of carbohydrate reserves in oak and beech are discussed with reference to differences in phenology, early spring growth and hydraulic properties between ring-porous trees and diffuse-porous trees.