Cell wall dimensions reign supreme: cell wall composition is irrelevant for the temperature signal of latewood density/blue intensity in Scots pine

Many microdensitometric techniques are available for deriving maximum latewood density (MXD), which is the state-of-the-art proxy parameter for local to hemispheric-scale temperature reconstructions of the last millennium. Techniques based on X-ray radiation and visible light reflection, such as “blue intensity” (BI), integrate both the density/composition and the dimensions of the cell walls to derive microdensitometric data. In contrast, the dendroanatomical technique relies only on the dimensions of the cell walls. It is therefore possible to isolate cell wall variables by subtracting data derived using the dendroanatomical technique from data derived using X-ray and BI-based techniques.In this study, we explore differences in well-replicated data from parallel X-ray, BI, and dendroanatomical measurements of temperature-sensitive Pinus sylvestris trees from northern Finland. We aim to determine whether cell wall density is critical to the success of X-ray-based MXD, and whether the BI-based parameter counterpart, here termed MXBI, contains useful information about the composition of the cell wall (specifically the lignin).Our results indicate that cell wall density and cell wall BI have no relevant influence on MXD and MXBI measurements. Even in years with severely reduced lignification, identified as so-called “blue rings”, dendroanatomical MXD (aMXD) measurements do not deviate significantly from their MXD or MXBI counterparts. Moreover, derived chronologies of cell wall density and cell wall BI contain no significant climate signals when correlated with local climate. Maximum latewood density of conifers can thus be obtained without bias using the dendroanatomical technique. Because lignin content appears to play a negligible role for cell wall BI, the cell wall BI likely presents the biggest challenge when producing unbiased MXBI data. This is because BI data is notorious for cell wall color distortion across the heartwood and sapwood, and between living wood and dead wood, and may therefore distort the otherwise strong link with wood density on multidecadal scales.     

xRing—An R package to identify and measure tree-ring features using X-ray microdensity profiles

Climate influences tree-ring density and ring-density variables extracted from X-ray images have been widely used for climate reconstructions. The R package xRing was developed to identify and measure tree rings on X-ray microdensity profiles automatically. This package is available for free and it offers functions to visualize and calibrate X-ray images, to detect tree-ring borders and to identify earlywood-latewood transition using wood density variations at the inter- and the intra-ring scale. The most important functions are calibrateFilm, detectRings, correctRings, detectEwLw, and getDensity. Outputs of these functions are S3 objects, for which specific methods are provided, including plot and print. The non-linear relationship between optical density of the film and wood density is defined by the function calibrateFilm. The function detectRings detects tree rings using wood density profiles as input. This function uses the difference between local maximum and minimum values to identify tree-ring borders automatically. The correctRings function is used to call a Graphical User Interface (GUI) to visualize and to correct tree-ring borders manually. After correcting tree-ring borders, the detectEwLw function is used to compute earlywood and latewood widths by dividing rings according to relative intra-ring density changes. The getDensity function computes for each tree ring the minimum (maximum) density and the mean earlywood, latewood and whole-ring density. Finally, a list with dataframes with tree-ring width and density variables can be obtained using the function getRwls. One of the major advantages of xRing package is that requires little knowledge of R language, but at the same time it can be easily changed or adapted by experienced users.     

Relationship between wood anatomy,tree-ring widths and wood density of Pinus sylvestris L. and climate at high latitudes in northern Sweden

In this study, wood anatomy, tree-ring width and wood density of Pinus sylvestris at the northern timberline in Fennoscandia were used to identify relationships among the parameters and to screen them for their climatic signals. Furthermore we investigated the influence of the juvenile wood section for all parameters developed. The measurements of wood anatomy were conducted with confocal laser scanning microscopy (CLSM) while the density profiles were produced using an Itrax MultiScanner. We developed chronologies of ring width, wood density and anatomy for a period between 1940 and 2010. Correlations between wood density and wood anatomy were strong in the latewood part. For some wood anatomy and density chronologies youth trends were found in the juvenile part. Wood density decreased from the pith up to the 9th ring and stabilized afterwards, while cell lumen diameter and lumen area increased simultaneously up to the 15th ring. All chronologies contained strong summer temperature signals. The wood anatomical variables provided additional information about seasonal precipitation which could not be found in wood density and tree-ring widths. Our study confirmed previous results stating that the parameter maximum density contains the strongest climate signal, that is, summer temperatures at the northern timberline. Nevertheless, the intra-annual data on tracheid dimensions showed good potential to supply seasonal climatic information and improve our understanding of climatic effects on tree growth and wood formation.     

Dendroclimatic potential of dendroanatomy in temperature-sensitive Pinus sylvestris

The most frequently and successfully used tree-ring parameters for the study of temperature variations are ring width and maximum latewood density (MXD). MXD is preferred over ring width due to a more prominent association with temperature. In this study we explore the dendroclimate potential of dendroanatomy based on the first truly well replicated dataset. Twenty-nine mature living Pinus sylvestris trees were sampled in North-eastern Finland at the cool and moist boreal forest zone, close to the latitudinal tree line, where ring width, X-ray MXD as well as the blue intensity counterpart MXBI were compared with dendroanatomical parameters. Maximum radial cell wall thickness as well as anatomical MXD and latewood density appeared to be the most promising parameters for temperature reconstruction. In fact, these parameters compare favorably to MXD derived from X-ray techniques as well as MXBI, in terms of shared variation and temperature correlations across frequencies and over time. The reasons for these results are thought to be the unprecedentedly high measurement resolution of the anatomical technique, which provide the optimal resolution – the cell – whereas X-ray techniques have a slightly lower resolution and BI techniques even lower. While the results of this study are encouraging, further tests on longer and multigenerational chronologies are required to more generally and fully assess the dendroclimate potential of anatomical parameters.     

Evaluation of X-ray densitometry to identify tree-ring boundaries of two deciduous species from semi-arid forests in Brazil

The presence of visible annual rings in semi-arid tropical trees may allow the application of dendrochronological methods. However, variation in water availability may cause the formation of narrow, irregular ill- or non-defined annual rings hindering the correct dating of tree-ring series. We aimed to evaluate X-ray densitometry as a method to identify tree rings of two deciduous tree species from the Caatinga forest, a semi-arid region in the northeast of Brazil, and compare with two other methods commonly used in dendrochronology, the sliding-stage micrometer and image analysis. Xylem was observed macro- and microscopically and wood anatomical features were assessed in Aspidosperma pyrifolium and Poincianella pyramidalis trees. In both species, tree-ring boundaries were identified considering intra-annual density patterns and wood anatomical features. No significant differences in tree-ring widths were found among methods. X-ray densitometry measurements showed a positive correlation with the measurements obtained with image analysis and sliding-stage micrometer in A. pyrifolium and P. pyramidalis, revealing the high reliability of the methods used. However, inter-correlation of tree-ring width series showed differences in the accuracy of crossdating across measuring methods. The maximum, mean and minimum density values were species-dependent, with mean wood density of A. pyrifolium lower than P. pyramidalis. Our results highlight X-ray densitometry as an important and complementary tool to identify tree-rings boundaries in semi-arid tree species, especially in A. pyrifolium. Along with other measuring methods, it may provide higher accuracy in dendrochronological studies in semi-arid or subtropical environments.     

黑龙江漠河兴安落叶松与樟子松树轮生长特性及其对气候的响应

树木年轮 (简称树轮 ) 气候学是监测与重建全球气候变化的重要方法之一。针叶树树轮的生长能反馈出气温的变化, 在高纬度地带尤为明显。该文分析了生长在我国最北部的兴安落叶松 (Larixgmelinii) 与樟子松 (Pinussylvestrisvar.mongolica) 的树轮密度和宽度的特性。落叶松最大密度、晚材平均密度、早晚材宽度和轮宽都远高于樟子松。樟子松的所有密度变量的样本方差都明显高于兴安落叶松, 宽度变量的样本方差却明显低于兴安落叶松。两树种密度变量的差值年表显著相关, 宽度变量之间没有显著相关关系。落叶松与樟子松的晚材密度的形成受 7、8月的最高温控制。另外, 樟子松的晚材还与生长季节的长短相关。落叶松的年轮宽度对生长季节开始前的温度敏感, 而樟子松的轮宽对气候变量没有很好的响应。结果表明, 落叶松与樟子松的树轮最大密度都与生长季后期的温度显著相关, 两树种的树轮信息对气候变化的重建有很大的潜力。     

Temperature sensitivity of blue intensity,maximum latewood density,and ring width data of living black spruce trees in the eastern Canadian taiga

The blue intensity (BI) technique provides opportunities to obtain surrogates to tree-ring density for reconstructing summer temperatures in high-latitude regions. In this study, we compare latewood BI (LBI) and delta BI (DBI), with the conventional X-ray maximum latewood density (MXD) and tree-ring width (TRW) data using 178 living trees of black spruce (Picea mariana (Mill.) B.S.P.), one of the most dominant species of conifers in the Northern Hemisphere, from 17 sites across the eastern Canadian taiga. The regional LBI and DBI chronologies are highly correlated to that of MXD (Pearson’s r = 0.97 and 0.92, respectively), while DBI is also similar to TRW (Pearson’s r = 0.67). Both LBI and DBI exhibit stronger responses to the May–August temperatures than TRW over larger time and spatial scales. However, only DBI is comparable to MXD data from inter-annual to decadal timescales. Low-frequency components of LBI data are likely distorted by color biases even if no obvious discoloration is present, as well as by the potentially low measurement resolution, leading to an overall weaker temperature sensitivity compared to the MXD data. Resampling experiments suggest that a minimum replication of 10 trees is needed to retain ≥90 % of the optimal temperature signal for MXD, LBI, and DBI data, and a minimum of 20 trees is required for TRW data.     

Blue intensity from a tropical conifer’s annual rings for climate reconstruction: An ecophysiological perspective

We developed Blue Intensity (BI) measurements from the crossdated ring sequences of Fokienia hodginsii (of the family Cupressaceae) from central Vietnam. BI has been utilized primarily as an indirect proxy measurement of latewood (LW) density of conifers (i.e., LWBI) from high latitude, temperature-limited boreal forests. As such, BI closely approximates maximum latewood density (MXD) measurements made from soft x-ray. The less commonly used earlywood (EW) BI (EWBI) represents the minimum density of EW and is influenced by the lighter pixels from the vacuoles or lumens of cells. The correlation of our BI measurements with climate, strongest for EWBI, rivals that for total ring width (RW), and we demonstrate that it can be successfully employed as an independent predictor for reconstruction models. EWBI exhibits robust spatial correlations with winter and spring land temperature, sea surface temperature (SST) over the regional domain of ENSO, and the Standardized Precipitation Evapotranspiration Index (SPEI) over Indochina. However, in order to mitigate the effects of color changes at the heartwood – sapwood boundary we calculated ΔBI (EWBI-LWBI), and it too exhibits a significant (p < 0.05), temporally stable response to prior autumn (Oct-Nov) rainfall and winter (December to April) dry season temperature. We interpret this response as reflecting a potential cavitation defense by reducing lumen diameter as a means to safeguard hydraulic conductivity in the stem, and to prevent the xylem from imploding due to negative pressure. This study has wide implications for the further use of BI from the global tropics, though it is unclear how many tropical tree species will be appropriate for use. It seems very likely that other wood anatomical measurements can be combined with BI and RW for climate reconstruction.     

X-ray microdensitometry applied to subfossil tree-rings: growth characteristics of ancient pines from the southern boreal forest zone in Finland at intra-annual to centennial time-scales

A collection of subfossil wood of Pinus sylvestris (Scots pine) was exposed to X-ray densitometry. The collection of 64 samples from the southern boreal forest zone was dendrochronologically cross-dated to a.d. 673-1788. Growth characteristics were determined by performing density profiles including the following parameters: minimum density, earlywood and latewood boundary density, maximum density, earlywood width, earlywood density, latewood width, latewood density, annual ring width and annual ring density. Seven out of the nine parameters were found to contain non-climatic growth trends and six were found to be heteroscedastic in their variance. Tree-specific records were indexed, to remove the non-climatic growth trends and stabilize the variance, and combined into nine parameter-specific tree-ring chronologies. Growth characteristics of the pines changed in parallel with the generally agreed climatic cooling from the Medieval Warm Period to the Little Ice Age: pine tree-rings showed decreasing maximum densities from the period a.d. 975-1150 to a.d. 1450–1625. A concomitant change in the intra-annual growth characteristics was detected between these periods. The findings indicate that not only the trees growing near the species’ distributional limits are sensitive to large-scale climatic variations but also the trees growing in habitats remote from the timberline have noticeably responded to past climate changes.     

Wood properties and ring width responses to long-term atmospheric CO2 enrichment in field-grown loblolly pine (Pinus taeda L.)

Loblolly pine (Pinus taeda L.) were grown in the field, under non-limiting nutrient conditions, in open-top chambers for 4 years at ambient CO₂ partial pressures (pCO₂) and with a CO₂-enriched atmosphere (+ 30 Pa pCO₂ compared to ambient concentration). A third replicate of trees were grown without chambers at ambient pCO₂. Wood anatomy, wood density and tree ring width were analysed using stem wood samples. No significant differences were observed in the cell wall to cell lumen ratio within the latewood of the third growth ring formed in 1994. No significant differences were observed in the density of resin canals or in the ratio of resin canal cross-sectional area to xylem area within the same growth ring. Ring widths were significantly wider in the CO₂-enrichment treatment for 3 of 4 years compared to the ambient chamber control treatment. Latewood in the 1995 growth ring was significantly wider than that in the ambient control and represented a larger percentage of the total growth-ring width. Carbon dioxide enrichment also significantly increased the total wood specific gravity (determined by displacement). However, when determined as total sample wood density by X-ray densitometry, the density of enriched samples was not significantly higher than that of the ambient chamber controls. Only the 1993 growth ring of enriched trees had a significantly higher maximum latewood density than that of trees grown on non-chambered plots or ambient chambered controls. No significant differences were observed in the minimum earlywood density of individual growth rings between chambered treatments. These results show that the most significant effect of CO₂ enrichment on wood production in loblolly pine is its influence on radial growth, measured as annual tree ring widths. This influence is most pronounced in the first year of growth and decreases with age.     

The importance of early summer temperature and date of snow melt for tree growth in the Siberian Subarctic

Wood material for at least 12 larch trees at six sites [Larix sibirica Ldb, Larix gmelinii (Rupr.) Rupr, Larix cajanderi Mayr] near the northern timberline in Siberia was analyzed to investigate influence of climatic factor changes on tree-ring growth at high latitudes. Tree-ring cell size, maximum latewood density and ring width measured by means of image analysis and X-ray radiodensitometry and calculated latewood cell-wall thickness were used. Correlation analysis of tree-ring structure parameter chronologies with temperatures averaged over periods of 5 days (pentad) shows that early summer temperature (mean for 5-6 pentads, depending on the region, starting from the middle of June) and date of snow melt are the most important factors that define seasonal growth and tree-ring structure. Analysis of instrumental climatic data indicates that a positive trend of early summer temperature was combined with winter precipitation (October-April) increase and this combination leads to later snow melt. Based of the results of tree-ring growth modelling, it was shown that later snow melt (hence, delayed initiation of cambial activity and, as a result, decrease of wood production) explains the changes in the relationship between tree ring width and summer temperature dynamics observed after the 1960s for a large area of the Siberian Subarctic. The understanding of the role of winter precipitation in controlling ring growth, through its effect on the timing of cambial activation, suggests the possibility of using ring structure parameters to create reconstructions of past winter precipitation variations.     

Dendrochronology of maritime pine in the middle of the Atlantic Ocean

The Azores Archipelago, located in the North Atlantic Ridge, experiences heavy rainfall and mild temperatures with weak seasonal differences due to oceanic influence. To our knowledge, there have been no dendrochronological studies in the Azores. The aim of this study is to explore the dendrochronological potential of Pinus pinaster Ait. growing in this archipelago and to determine what limiting factor is regulating tree growth. To do so, we have sampled adult maritime pine trees growing in a plantation, in the Pico island of the Azores.Tree ring boundaries were not always easily distinguished, suggesting that in some years cambial activity did not stop during winter. Despite this, it was possible to successfully crossdate the tree-ring series and to establish a tree-ring width chronology with a strong common signal. Climatic correlations revealed a positive response to spring precipitation but no temperature signal in the tree-ring width chronology. Tree-ring width was also negatively correlated with the North Atlantic Oscillation (NAO) and the sea level pressure (SLP) in May − June.Intra-annual density fluctuations (IADFs), which are anatomical features formed in response to variations in environmental conditions during the growing season, were present in 85% of the tree rings. IADFs were identified based on its position within the ring: type E⁺, characterized as a transition wood from early- to latewood; type L, the most frequent, characterized as earlywood-like cells within latewood; and type L⁺, characterized as earlywood-like cells between latewood and earlywood of the next tree ring. Each IADF type presented a unique climatic signal: type E⁺ was positively correlated with early summer precipitation and early spring temperature; type L was positively correlated with early autumn precipitation and temperature; and type L⁺ was positively correlated with late autumn precipitation.In conclusion, the tree-ring width chronology established for maritime pine growing in the Pico Island of Azores contains a clear climatic signal for spring precipitation, whereas IADFs frequency correlated better with precipitation later in the growing season. For this reason, we suggest that IADFs should be included in future dendrochronological studies in the Macaronesia Biogeographical region since they can improve the climatic signal present in tree-ring width chronologies.     

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。     

The use of X-ray densitometry to evaluate the wood density profile of Tectona grandis trees growing in fast-growth plantations

Tectona grandis (teak) is an important commercial tree species that is widely used in tropical dendrochronology due to the formation of climate-sensitive annual growth rings. However, young trees growing in plantation conditions exhibit poor ring visibility during the first years of growth, limiting the dendrochronology application. In the present study, we use x-ray densitometry to determine the wood density profile between and within annual rings and at the sapwood-heartwood boundary in trees from fast-growth plantations. The resulting wood density profiles (WDP) can be categorized as uniform, stable growth, unstable growth, and false. The annual ring boundaries were indistinct in trees less than 8 years old. In mature trees, the annual ring boundaries are more defined. In relation to the sapwood-heartwood boundary, the WDP showed a decrease in the wood density; however, this decrease is influenced by the annual ring boundary when the two boundaries coincide. The identification of annual rings in trees growing in fast-growth plantations should be combined with X-ray densitometry and visual identification if wood density data are necessary for deriving other analysis, as climate change, from annual ring.     

Dendroclimatic signals deduced from riparian versus upland forest interior pines in North Karelia, Finland

Radial growth of boreal tree species is only rarely studied in riparian habitats. Here we investigated chronologies of earlywood, latewood, and annual ring widths and blue intensity (BI; a surrogate to latewood density) from riparian lake shore and upland forest interior pines (Pinus sylvestris L.) growing in boreal forest in eastern Finland. Riparian and upland chronologies were compared to examine differences in the pine growth variability and growth response to climatic variation in the two habitats. It was found that the climatic variables showing statistically significant correlations with the tree-ring chronologies were related to snow conditions at the start of the growing season. Deeper snowpack led to reduced upland pine growth, possibly due to delayed snowmelt and thus postponed onset of the growing season. Warm late winters were followed by increased riparian pine growth because of earlier start of the snow-melt season and thus a lower maximum early summer lake level. Moreover, riparian pines reacted negatively to increased rainfall in June, whereas the upland pines showed a positive response. Latewood growth reacted significantly to summer temperatures. The BI chronology showed a strong correlation with warm-season temperatures, indicating an encouraging possibility of summer temperature reconstruction using middle/south boreal pine tree-ring archives.     

Yearly maps of summer temperatures in Western Europe from A.D. 1750 to 1975 and Western North America from 1600 to 1982:

The primary purpose of this publication is to make available two series of historical maps showing summer (April–September mean) temperatures over western Europe from A.D. 1750 and western North America from A.D. 1600. These maps have been reconstructed using networks of temperature-sensitive tree-ring chronologies as part of a continuing collaboration between the Swiss Federal Institute for Forest, Snow and Landscape Research (FSL) at Birmensdorf, Switzerland and the Climatic Research Unit (CRU) at the University of East Anglia, Norwich, U.K. The maps were reconstructed using relatively long maximum latewood density chronologies selected from the extensive networks of tree-ring data produced at the FSL. The reconstructions were produced using a spatial regression technique developed at CRU.The basis for the reconstructions are wood samples from various conifers growing at cool-moist sites close to the alpine or boreal timberlines. This material was analysed by x-ray densitometry and the resulting maximum latewood density chronologies were calibrated against meteorological data over a 100-year calibration period.These results emphasize the value of dendroclimatological research which can provide continentalscale reconstructions of past climate which represent a potential resource for climatologists, biologists, and historians with an interest in the spatial variability of summer temperatures over the middle to high latitudes.     

Exploring wood anatomy,density and chemistry profiles to understand the tree-ring formation in Amazonian tree species

Long-term analysis of tree growth using annual tree rings is increasingly in demand for tropical tree species. The basis of these studies has traditionally been the anatomical identification of the annual ring boundary. However, the structure of these annual rings has been sparsely explored for complementary physical and chemical wood traits. Here, we explore the relationships among wood density features and chemical elements (S, K, Ca, Mn) involved in the annual tree ring formation of 12 tropical tree species from non-flooded forest in the southern Amazon basin. Transverse wood sections were used for each species to determine: 1) macroscopic distinction (radial growth and wood density), 2) microscopic analyse of vessels, axial and ray parenchyma (anatomy) and 3) X-ray densitometry (physical) and X-ray fluorescence (chemical). For some species, the profiles of wood density, and Ca and Mn content showed intra- and inter-annual patterns that allowed to define and characterize the growth boundary of tree rings. Ca, K and S were mainly distributed in axial parenchyma cells, and around vessels, whereas, Mn was mainly distributed in fibres. Our results showed significant species-specific correlations between tree-ring width, density and concentrations of Ca, K and Mn. The anatomical characterization and the complementary information provided by the density and chemical profiles in some Amazonian species can represent a valuable proxy to improve the definition of annual ring-boundaries and improve the understanding of long-term growth and physiological patterns.     

Comparison of methods for the demarcation between earlywood and latewood in tree rings of Norway spruce

The precise demarcation between earlywood and latewood is important for the detailed analysis of intra-annual tree ring features. Different techniques based on visual assessment, wood anatomy analysis and X-ray densitometry have been developed and are currently used for this purpose. Depending on the chosen method, tree species and environmental conditions, the results can significantly vary. Thus, it is important to determine the technique optimal for a particular research. Here, we investigated Norway spruce (Picea abies) tree rings to examine the agreement among the following demarcation methods: (1) direct visual assessment, (2) Mork’s index (anatomical definition of the transition from earlywood to latewood based on cell wall-lumen ratio) and (3) fixed and floating density thresholds applied to intra-ring density profiles. The aim was to modify both the Mork’s criterion and density thresholds on the basis of reference values given by visual identification of earlywood/latewood transition. A total of 231 tree rings were analysed by all methods. Our results showed that the usage of floating threshold (defined for each ring separately based on density profiles) is more reliable in comparison with fixed threshold (the same threshold value used for all tree rings and samples). Statistical analysis revealed the best correspondence between visual identification of earlywood/latewood transition and demarcation based on the standard Mork’s index and the floating density threshold derived as 80 % of maximum latewood density. In terms of Mork’s index calibration, the results showed that to determine latewood cells in Norway spruce trees growing in temperate conditions, it is sufficient to use an index value equal to 0.83. The results are applicable for the studied spruce population growing in a temperate climate. The methodology itself, however, is universal and can help to calibrate criteria for earlywood-latewood demarcation under specific conditions.     

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.     

The potential of intra-annual density information for crossdating of short tree-ring series

The crossdating of tree-ring series is typically based on tree-ring width sequences, which is a crude abstraction of the growth signal stored in tree rings. In contrast, intra-annual wood density data allows a much more detailed comparison of wood growth processes and new measurement techniques scale well to measure large amounts of samples. Thus, chronologies of intra-annual densitometric curves can be built. Here, we investigate to what extent intra-annual wood density information can improve crossdating. We evaluate different approaches on a data set of Norway spruce trees (Picea abies) and compare the results to standard methods that are based on ring width or maximum density. Our results show that intra-annual densitometric data indeed increases crossdating success rate notably for short tree ring series that cover less than 25 years.