Radial-growth and wood-anatomical changes in overaged Quercus pyrenaica coppice stands: functional responses in a new Mediterranean landscape

Recent land-use changes in intensively managed forests such as Mediterranean coppice stands might profoundly alter their structure and function. We assessed how the abandonment of traditional management practices in coppice stands, which consisted of short cutting-cycles (10–15 years), has caused overaging (stems are usually much older than when they were coppiced) and altered their wood anatomy and hydraulic architecture. We studied the recent changes of wood anatomy, radial growth, and hydraulic architecture in two stands of Quercus pyrenaica, a transitional Mediterranean oak with ring-porous wood forming coppice stands in W–NW Spain. We selected a xeric and a mesic site because of their contrasting climates and disturbance histories. The xeric site experienced an intense defoliation after the severe 1993–1994 summer drought. The mesic site was thinned in late 1994. We studied the temporal variability in width, vessel number and diameter, and predicted the hydraulic conductivities (K _h) of earlywood and latewood. In the mesic site, we estimated the vulnerability to xylem cavitation of earlywood vessels. Overaging caused a steep decline in latewood production at a cambial age of 14 years., which was close to the customary cutting cycle of Q. pyrenaica. The diameter distribution of vessels was bimodal, and latewood vessels only accounted for 4% of the K _h. Overaging, acting as a predisposing factor in the decline episode, was observed at the xeric site, where most trees did not produce latewood in 1993–1995. At the mesic site, thinned trees formed wider tree-rings, more latewood and multiseriate tree-rings than overaged trees. The growth enhancement remained 8 years after thinning. Most of the hydraulic conductivity in earlywood was lost in a narrow range of potentials, between −2.5 and −3.5 MPa. We have shown how hydraulic conductivity and radial growth are closely related in Q. pyrenaica and how aging modulates this relationship.     

Wood borer performance and wood characteristics of drought‐stressed Scots pine seedlings

Four‐year‐old Scots pine [Pinus sylvestris L. (Pinaceae)] seedlings were exposed to medium and severe drought stress for two consecutive years. The anatomical properties of drought‐stressed Scots pine wood and their impact on the performance of destructive wood boring early instars of Hylotrupes bajulus L. (Coleoptera: Cerambycidae) were studied. Drought stress significantly decreased diameter of earlywood tracheids in both growing years and diameter of latewood tracheids after the second growing season only. Cell lumen area was significantly decreased by both medium and severe drought stress compared to well‐watered controls. In addition, area of cell lumen was significantly smaller in severe drought than in medium drought treatment. The drought stress marginally increased the number of resin canals in the wood, but did not affect the size of resin canals either in wood or bark. The relative growth rate of xylophagous H. bajulus neonatal larvae was not significantly affected by drought stress during the 106‐day feeding period on Scots pine wood blocks. The results show that although water availability was an important factor affecting the development and anatomy of wood cells, observed changes in wood characteristics did not affect the performance of early instars feeding on wood processed from drought‐stressed young Scots pine seedlings.     

太行山南麓不同径级栓皮栎生长对气候要素及干旱事件的响应

为评估径级对树木的气候-生长关系的影响,建立太行山南麓低海拔地区栓皮栎全轮、早材、晚材宽度年表,对比两个径级栓皮栎人工林径向生长对气候响应的敏感性差异,使用叠加时代分析揭示干旱事件对不同径级栓皮栎的影响,为气候变化背景下研究区栓皮栎人工林可持续经营提供参考数据。结果表明: 大径级栓皮栎全轮、晚材年表的平均敏感度高于小径级栓皮栎年表,但小径级栓皮栎早材年表的平均敏感度高于大径级栓皮栎早材年表。晚材是树轮气候响应最敏感的组分。两径级栓皮栎标准年表指数对当年气候因子的响应模式相似。小径级栓皮栎全轮、晚材生长对当年6—8月的气候因子更敏感,其早材对生长季之前(1—2月)气候因子更敏感;而大径级栓皮栎早材对去年的气候因子更敏感,有更强的滞后效应。研究区栓皮栎对干旱非常敏感,干旱年份两径级栓皮栎全轮、晚材的径向生长量显著低于上一年,且大径级栓皮栎的减少幅度更高,但干旱后大径级栓皮栎生长恢复程度更高。     

Intra-annual wood anatomical features of high-elevation conifers in the Great Basin,USA

Mountain conifers in the Great Basin of North America have provided some of the longest, continuous, and annually resolved paleoclimate records. Climate-growth relationships at the cellular level, which help understand wood formation processes that underlie dendroclimatic reconstructions, are at present largely unexplored in the Great Basin. We analyzed 42 trees located in the Snake Range (eastern Nevada, USA) at three sites along an elevation gradient. Sampled species included white fir (Abies concolor), Douglas fir (Pseudotsuga menziesii), limber pine (Pinus flexilis), bristlecone pine (Pinus longaeva), and Engelmann spruce (Picea engelmannii). Wood anatomical features were quantified for two consecutive years, 2011 and 2012. Lumen area, cell wall thickness, lumen diameter, and wall-to-cell ratio were measured for the total ring as well as for earlywood and latewood. Mean standardized tracheidograms highlighted differences between 2011 and 2012, in particular concerning lumen area and wall-to-cell ratio. Most annual variation was due to earlywood, rather than latewood. Anatomical parameters of limber pine, the only species that could be tested at both the montane and subalpine sites, varied with elevation. Principal component analysis showed that the main axis of variability was related to dimensional parameters (e.g. lumen area), which reflected differences in water availability.     

Size-dependent responses to summer drought in Scots pine, Norway spruce and common oak

In this study, we provide a detailed analysis of tree growth and water status in relation to climate of three major species of forest trees in lower regions of Bavaria, Southern Germany: Scots pine (Pinus sylvestris), Norway spruce (Picea abies) and common oak (Quercus robur). Tree-ring chronologies and latewood δ¹³C were used to derive measures for drought reaction across trees of different dimensions: growth reduction associated with drought years, long-term growth/climate relations and stomatal control on photosynthesis. For Scots pine, growth/climate relations indicated a stronger limitation of radial growth by high summer temperatures and low summer precipitation in smaller trees in contrast to larger trees. This is corroborated by a stronger stomatal control on photosynthesis for smaller pine trees under average conditions. In dry years, however, larger pine trees exhibited stronger growth reductions. For Norway spruce, a significantly stronger correlation of tree-ring width with summer temperatures and summer precipitation was found for larger trees. Additionally, for Norway spruce there is evidence for a change in competition mode from size-asymmetric competition under conditions with sufficient soil water supply to a more size-symmetric competition under dry conditions. Smaller oak trees showed a weaker stomatal control on photosynthesis under both dry and average conditions, which is also reflected by a significantly faster recovery of tree-ring growth after extreme drought events in smaller oak trees. The observed patterns are discussed in the context of the limitation-caused matter partitioning hypothesis and possible species-specific ontogenetic modifications.     

Driving factors of a vegetation shift from Scots pine to pubescent oak in dry Alpine forests

An increasing number of studies have reported on forest declines and vegetation shifts triggered by drought. In the Swiss Rhone valley (Valais), one of the driest inner‐Alpine regions, the species composition in low elevation forests is changing: The sub‐boreal Scots pine (Pinus sylvestris L.) dominating the dry forests is showing high mortality rates. Concurrently the sub‐Mediterranean pubescent oak (Quercus pubescens Willd.) has locally increased in abundance. However, it remains unclear whether this local change in species composition is part of a larger‐scale vegetation shift. To study variability in mortality and regeneration in these dry forests we analysed data from the Swiss national forest inventory (NFI) on a regular grid between 1983 and 2003, and combined it with annual mortality data from a monitoring site. Pine mortality was found to be highest at low elevation (below 1000 m a.s.l.). Annual variation in pine mortality was correlated with a drought index computed for the summer months prior to observed tree death. A generalized linear mixed‐effects model indicated for the NFI data increased pine mortality on dryer sites with high stand competition, particularly for small‐diameter trees. Pine regeneration was low in comparison to its occurrence in the overstorey, whereas oak regeneration was comparably abundant. Although both species regenerated well at dry sites, pine regeneration was favoured at cooler sites at higher altitude and oak regeneration was more frequent at warmer sites, indicating a higher adaptation potential of oaks under future warming. Our results thus suggest that an extended shift in species composition is actually occurring in the pine forests in the Valais. The main driving factors are found to be climatic variability, particularly drought, and variability in stand structure and topography. Thus, pine forests at low elevations are developing into oak forests with unknown consequences for these ecosystems and their goods and services.     

Intra-annual tree-ring parameters indicating differences in drought stress of Pinus sylvestris forests within the Erico-Pinion in the Valais (Switzerland)

The effects of drought on radial growth of Pinussylvestris were investigated by comparing sites along hydricgradients. The gradients were located in Valais, an inner Alpine dry valley inSwitzerland, with each consisting of two site types, an extreme dry, xeric siteand a less dry, moderate site. The two site types were assigned tophytosociological associations within the Erico-Pinion. The investigationcovered the responses of tree growth to climate and particularly concentratedonintra-annual features of tree-rings such as earlywood/latewood ratio,intra-annual density fluctuations (IADFs) and traumatic tissues (TTs) as wellasthe sapwood/heartwood ratio. Radial growth differed according to the sitetypes,with trees on dry sites generally showing more missing rings, lower mean ringwidths, lower autocorrelation, higher mean sensitivities, reduced latewoodproportions and lower sapwood areas than trees on moderate sites. Therelationships between climate and tree-ring width, studied using responsefunction analysis, varied strongly between the site types within theErico-Pinion: Tree growth on dry sites was positively influenced byprecipitation at the end of the winter and the beginning of the growing seasonand negatively influenced by temperature in June. Winter precipitation waspositively correlated with radial growth, demonstrating its importance for thesuccessful root and shoot growth of the plants in spring on dry sites. Onmoderate sites, tree growth was less controlled by climate than by priorgrowth.The intra-annual density fluctuations (IADFs) provided a valuable means todifferentiate between the site types. In comparison to the moderate sites, thetrees on dry sites contained more IADFs, and their frequency was increased.Moist-cool conditions in the middle of the growing season were the triggeringfactor for IADFs on dry sites, whereas on moderate sites, there must be anadditional warm period in early summer in order to initiate IADFs. Most IADFswere found in latewood. We found no relationship between climate and traumatictissues (TTs). It is unclear whether other abiotic or biotic factors such aswounding by insects or birds are responsible for the development of TTs. Theassignment of these differences in tree growth behaviour to phytosociologicalassociations will enable a deeper understanding of the site types and willfacilitate the comparison with similar studies. Furthermore, the results can becombined with studies from other scientific disciplines concerning thesephytosociological associations. The ecological indicator values of thevegetation was a precise method for the distinction of site types.     

Long-term radial growth and climate-growth relationships of Quercus petraea (Matt.) Liebl. and Quercus cerris L. in a xeric low elevation site from Hungary

Sessile oak (Quercus petraea [Matt.] Liebl.) and Turkey oak (Quercus cerris L.) dominated mixed forests are common in low montane and hilly regions in Hungary. Here, we aimed to describe the long-term pattern and climatic responses of the radial growth of Q. petraea and Q. cerris in a xeric low-elevation forest, using retrospective tree-ring analysis for the period 1910–2019. We performed separate analyses with time series of full tree-ring (TRW), earlywood (EW) and latewood (LW) widths. Our results showed that the radial growth of the two species was largely synchronous over time, but became transiently divergent for a 20-year period after a drought in 1968, due to the greater-than-expected growth of Q. cerris and the supressed growth of Q. petraea. Precipitation was the major growth-limiting factor for both species, with a strong positive influence on LW particularly during the current early growing season (March-June), on EW in the previous late summer (August-September) and in the current early spring (March), and on all tree-ring traits in the previous December. The radial growth of both species was negatively related to temperature in the spring (May) and late summer (August) of both current and previous years. The climate-growth relationships showed general instability over time: the most striking temporal change was a gradual shift of positive correlations with precipitation and SPEI during the growing season from spring (March-May) to summer (June-August) since the 1980s over the analysed period. The two species had similarly low growth resistance to droughts in four studied pointer years (1968, 1993, 2002 and 2012), but Q. cerris exhibited a greater capacity to recover over the four post-drought years, and thus higher growth resilience, particularly after the drought of 1968. Our results contribute to the better understanding of the role of climate variability and droughts in the growth of the two co-existing species in transitional locations between closed forests and forest-steppes.     

Long-term changes in wood density and radial growth of Quercus petraea Liebl. in northern France since the middle of the nineteenth century

Long-term changes in sessile oak (Quercus petraea Liebl.) growth and wood density were studied using cores collected from 99 even-aged high forest stands between 56 and 187 years old, located in northeastern and north-central France. Growth and density trends were tested by analysis of variance and covariance. Two models were applied to two samples, sample A and sample B (sample B being a sub-sample with limited cambial age and calendar date ranges). Model 1 showed a significant increase in radial growth: +35%, +87% and +66% in earlywood width, latewood width and ring width, respectively, from 1811 to 1993 for sample A. Consequently, there was a positive trend in latewood ratio (+14%). A slight decrease in wood density was found: -3.3% and -5.4% for earlywood and latewood density, respectively. Despite an increase in latewood percentage, mean ring density showed a -2.0% decrease. Model 1 applied to a biomass indicator (density2ring width) showed a 62% increase from 10.4 to 16.8 kg m^-3 between 1811 and 1993 for sample A. Results for sample B were slightly different: the increase in latewood ratio was not detected. Model 2 showed a change with time in the positive hyperbolic relationship between mean density and ring width. The results are discussed. The decrease in wood density cannot be explained by N atmospheric deposition or by long-term changes in average temperature. Increasing atmospheric CO₂ levels cannot be invoked owing to the present lack of studies. Finally, hypotheses concerning long-term changes in wood anatomical characteristics are proposed.     

Climate–tree growth relationships of longleaf pine (Pinus palustris Mill.) in the Southeastern Coastal Plain, USA

Knowledge of tree growth/climate response relationships is important to dendroecological studies and dendroclimatic reconstructions, particularly in the Southeastern Coastal Plain where few such studies have been attempted. To this end, we developed tree-ring chronologies of total ring width, earlywood width, and latewood width from longleaf pine (Pinus palustris Mill.) at three sites in the Southeastern Coastal Plain to examine the climate–growth relationships for this tree species. The length of these chronologies is unprecedented for southern pine chronologies in the Southeast. We compared the tree-ring chronologies to monthly temperature, precipitation, Palmer drought severity index (PDSI), and Palmer hydrological drought index (PHDI) data from the pertinent climate divisions. We found that PDSI and PHDI have the highest correlation with longleaf pine growth, and the strongest relationships between longleaf pine growth and these variables occur between July and November. Precipitation in the spring and summer was also positively related to growth at all sites. The relationship between temperature and growth was the weakest among all climate variables, but warm summer temperatures had a consistent, negative relationship with longleaf pine growth. The climate signal in the latewood was generally more robust than for total ring width and earlywood width.     

Sensitivity of French temperate coniferous forests to climate variability and extreme events (Abies alba,Picea abies and Pinus sylvestris)

Questions: (1) How do extreme climatic events and climate variability influence radial growth of conifers (silver fir, Norway spruce, Scots pine)? (2) How do elevation and soil water capacity (SWC) modulate sensitivity to climate? Location: The sampled conifer stands are in France, in western lowland and mountain forests, at elevations from 400 to 1700 m, and an SWC from 50 to 190 mm. Methods: We established stand chronologies for total ring width, earlywood and latewood width for the 33 studied stands (985 trees in total). Responses to climate were analysed using pointer years and bootstrapped response functions. Principal component analysis was applied to pointer years and response function coefficients in order to elucidate the ecological structure of the studied stands. Results: Extreme winter frosts are responsible for greater growth reductions in silver fir than in Norway spruce, especially at the upper elevation, while Scots pine was the least sensitive species. Exceptional spring droughts caused a notable growth decrease, especially when local conditions were dry (altitude<1000 m and SWC<100 mm for silver fir, western lowlands for Scots pine). Earlywood of silver fir depended on previous September and November and current‐year February temperature, after which current June and July water supply influenced latewood. Earlywood of Norway spruce was influenced by previous September temperature, after which current spring and summer droughts influenced both ring components. In Scots pine, earlywood and latewood depended on the current summer water balance. Local conditions mainly modulated latewood formation. Conclusions: If the climate becomes drier, low‐elevation dry stands or trees growing in western lowlands may face problems, as their growth is highly dependent on soil moisture availability.     

Drought response of five conifer species under contrasting water availability suggests high vulnerability of Norway spruce and European larch

The ability of tree species to cope with anticipated decrease in water availability is still poorly understood. We evaluated the potential of Norway spruce, Scots pine, European larch, black pine, and Douglas‐fir to withstand drought in a drier future climate by analyzing their past growth and physiological responses at a xeric and a mesic site in Central Europe using dendroecological methods. Earlywood, latewood, and total ring width, as well as the δ¹³C and δ¹⁸O in early‐ and latewood were measured and statistically related to a multiscalar soil water deficit index from 1961 to 2009. At the xeric site, δ¹³C values of all species were strongly linked to water deficits that lasted longer than 11 months, indicating a long‐term cumulative effect on the carbon pool. Trees at the xeric site were particularly sensitive to soil water recharge in the preceding autumn and early spring. The native species European larch and Norway spruce, growing close to their dry distribution limit at the xeric site, were found to be the most vulnerable species to soil water deficits. At the mesic site, summer water availability was critical for all species, whereas water availability prior to the growing season was less important. Trees at the mesic were more vulnerable to water deficits of shorter duration than the xeric site. We conclude that if summers become drier, trees growing on mesic sites will undergo significant growth reductions, whereas at their dry distribution limit in the Alps, tree growth of the highly sensitive spruce and larch may collapse, likely inducing dieback and compromising the provision of ecosystem services. However, the magnitude of these changes will be mediated strongly by soil water recharge in winter and thus water availability at the beginning of the growing season.     

Are climatic factors responsible for the process of oak decline in Poland?

Oak decline, a complex process leading to increased mortality of this species, has been observed in Europe for many years. Previous studies suggest that climate conditions, especially drought, may be one of the most important factors that trigger this phenomenon. The paper investigates the radial growth and wood anatomy features of pedunculate oak (Quercus robur) trees of various health status as well as their response to climate conditions. Wood samples including all annual increments were taken at two sites (western and central Poland, 15 trees each). Based on the crown defoliation level, three health groups (healthy, weakened and dead oaks) were distinguished. Cross-sections were prepared with sliding microtome and Cell P image analysis software was used for the measurements. Tree-ring width (TRW), earlywood vessels density (VDen) and non-weighted vessels diameter (VD) were determined and correlated with mean monthly values of temperature, precipitation, vapour pressure, and Palmer Drought Severity Index (PDSI). Radial increment and anatomical parameters were significantly higher for the healthy oaks than for the weakened and the dead trees. TRW showed smaller dependence on climate than analysed anatomical attributes. No obvious pattern of relationship was found between oak radial growth and climate regarding tree health status. Our results revealed that the drought has a weak impact on the process of oak decline on investigated sites in Poland.     

Comparison of earlywood vessel variables in the wood of Quercus robur L. and Quercus petraea (Mattuschka) Liebl. growing at the same site

The differences in the microscopic structure of wood based on the variables of earlywood vessel area and tree ring width were analysed in 6 trees of Pedunculate oak (Quercus robur L.) and 6 trees of Sessile oak (Quercus petraea (Mattuschka) Liebl.) in the same forest stand at a site in the Vizovice Highland (Czech Republic). The aim of this paper was to assess any differences between the two oak species when grown in the same location. Also the effect of tree-ring width and the effect of age on earlywood vessel variables were analysed. Mean values of tree-ring widths were not different between species. Earlywood vessel area chronologies were synchronized well between species. The mean values of earlywood vessel area (average vessel area, average vessel area of the first row of vessels, area of the largest vessel) showed a significant difference. Finally, we can conclude that there were differences in vessel features between these oak species.     

A comparison of ring-width and event-year chronologies derived from white oak (Quercus alba) and northern red oak (Quercus rubra), southwestern Quebec, Canada

In dendrochronology, ring width has been a variable of choice when assessing the radial growth–climate association of tree species. We compared ring-width and event-year chronologies from a dendroclimatic perspective using both white oak (Quercus alba L.) and northern red oak (Quercus rubra L.). The study was conducted in three regions of the Ottawa valley in southern Québec. Twelve mixed red and white oak stands were selected and for each oak species, 12 chronologies were developed from tree-ring measurement and 12 others were derived using visual assessment of narrow or wide rings (event years). Ring-width and event-year chronologies gave almost identical results and revealed the prevalence of drought in the early growing season as the most influential factor in both species. This study emphasizes the utility of event-year chronologies in tree-ring studies and their comparativeness with ring width. Establishing event-year chronologies has the advantage of being faster than measuring ring width, it does not necessitate complex equipment and depending on the purpose of the study may prove to be at least comparable. The choice of species, their mean sensitivity, the ability to recognize narrow or large rings as well as the number of trees and sites to analyze may, however, be factors to consider when choosing to use event-year chronologies over the more commonly used ring-width ones.     

中国祁连山西段青海云杉树轮密度记录的气候信号

树轮气候学是研究过去气候变化的重要手段之一。以往研究表明,树轮密度是生长季温度的重要代用资料。本文建立了祁连山西段青海云杉132～135年的树轮最大密度年表、树轮最小密度年表、树轮晚材平均密度年表和树轮早材平均密度年表,比较了不同密度年表指示的最优气候信号及其季节组合,评估了其作为气候代用资料的潜力与价值。结果表明:树轮晚材平均密度和树轮早材平均密度对于树木生长季气候信号的响应远低于树轮最大密度和树轮最小密度;树轮早材平均密度和树轮最小密度与帕尔默干旱指数有很强的联系,表明树轮早材平均密度和树轮最小密度有成为干旱代用指标的潜力。     

Linkages between forest growth,climate, and agricultural production are revealed through analysis of seasonally-partitioned longleaf pine (Pinus palustris Mill.) tree rings

We established a five-century long tree-ring chronology partitioned between earlywood and latewood growth to examine intra-annual climate response and attempt to establish linkages to agricultural production. Longleaf pine earlywood and latewood width chronologies spanned the period 1491–2017 (527 years) and constitute one of the longest records achieved for this species. High monthly correlations were found between latewood growth and summer-fall Palmer Drought Z-Index. Correlations were consistently significantly positive for June through October. Intra-annual growth of earlywood and latewood were positively correlated for the full period of record, but exhibited variability in correlation strength through time. Conversely, earlywood and prior-year latewood were also frequently correlated, but correlations were found to switch between positive and negative association, possibly in response to Atlantic Ocean temperatures. Annual yields of major crops are coupled with latewood growth, representing a new and potentially valuable proxy for linking agricultural yields to climate proxies over multiple centuries.     

Anatomy of the vessel network within and between tree rings of Fraxinus lanuginosa (Oleaceae)

The three-dimensional (3-D) arrangement of vessels and the vessel-to-vessel connections in the secondary xylem of the stem of the ring-porous hardwood tree Fraxinus lanuginosa were studied in series of thick transverse sections with epifluorescence microscope and confocal laser scanning microscope. Vessels were traced in sequential sections, and vessel networks were reconstructed in two segments of wood with dimensions of 2 × 1.4 × 21.2 mm(3) and 2 × 1.4 × 5.8 mm(3) (tangential × radial × axial). The arrangement of vessels and intervessel pits were visualized by scanning electron microscopy in low-density polyethylene microcasts and on exposed tangential faces of growth-ring boundaries. The vessels deviated from the stem axis in the tangential direction and, to a lesser extent, in the radial direction. Some neighboring vessels were twisted around each other. Vessels that appeared solitary in single sections were found to be sequentially contiguous with a number of other vessels, forming networks that extended in the tangential direction and across growth-ring boundaries. In the 21.2-mm wood block, all earlywood vessels at the growth-ring boundary made contact with latewood vessels in the previous tree ring. Within a growth ring however, only a single contact was observed between individual earlywood and latewood vessels. Densely arranged intervessel pits were characteristic in the regions where adjacent vessels made contact with each other. Such bordered pits were abundant in the tangential walls of vessel elements adjacent to growth-ring boundaries. Therefore, bordered pits appear to provide the pathway for the radial transport of water via the vessel network across growth-ring borders. Fiber-tracheids, observed as terminal cells in the tree rings, might also contribute to the apoplastic transfer of water across ring borders.     

Drought as an Inciting Mortality Factor in Scots Pine Stands of the Valais, Switzerland

During the 20th century, high mortality rates of Scots pine (Pinus silvestris L.) have been observed over large areas in the Rhône valley (Valais, Switzerland) and in other dry valleys of the European Alps. In this study, we evaluated drought as a possible inciting factor of Scots pine decline in the Valais. Averaged tree-ring widths, standardized tree-ring series, and estimated annual mortality risks were related to a drought index. Correlations between drought indices and standardized tree-ring series from 11 sites showed a moderate association. Several drought years and drought periods could be detected since 1864 that coincided with decreased growth. Although single, extreme drought years had generally a short-term, reversible effect on tree growth, multi-year drought initiated prolonged growth decreases that increased a tree’s long-term risk of death. Tree death occurred generally several years or even decades after the drought. In conclusion, drought has a limiting effect on tree growth and acts as a bottleneck event in triggering Scots pine decline in the Valais.     

Effect of elevated [CO2] on stem wood properties of mature Norway spruce grown at different soil nutrient availability

The objective of the present study was to investigate the interactive effects of elevated [CO₂] and soil nutrient availability on secondary xylem structure and chemical composition of 41‐year‐old Norway spruce (Picea abies (L.) Karst.) trees. The nonfertilized and irrigated‐fertilized trees were, for 3 years, continuously exposed to elevated [CO₂] in whole‐tree chambers. Elevated [CO₂] decreased concentrations of soluble sugars, acid‐soluble lignin and nitrogen in stem wood, but the effects were not consistent between sampling height and/or fertilization. The effect of 2*ambient [CO₂] on wood structure depended on the exposure year and/or fertilization. Radial lumen diameter decreased and annual ring width increased in the second year of exposure (1999) in elevated [CO₂]. In the latter, the CO₂ effect was significant only in the nonfertilized trees. Stem wood chemistry and structure were significantly affected by fertilization. Fertilization increased the concentrations of nitrogen and gravimetric lignin, annual ring width, and radial lumen diameter. Fertilization decreased C/N ratio, mean ring density, earlywood density, latewood density, cell wall thickness, cell wall index, and latewood percentage. We conclude that elevated [CO₂] had only minor effects on wood properties while fertilization had more marked effects and thus may affect ecosystem processes and suitability of wood for different end‐use purposes.