Variation of different tree-ring parameters in samples from each terminal shoot of a Norway spruce tree

Variability of wood parameters in a tree is sometimes a rather nebulous concept since variability is evident within single cells, from early to latewood, from pith to bark and from stem base to the top of a tree. So far, stem analyses have been done using a restricted number of parameters, mostly ring-width, and using a restricted number of samples in the longitudinal direction. This study analyses a number of parameters from a single tree. An 81-year-old spruce tree was felled and internodial discs were taken from each annual terminal shoot. All tree rings in each disc were measured and a whole-stem analysis was completed for the following parameters: ring-width, mean ring density, maximum density, percentage of latewood, type of transition from early to latewood, intra-annual density fluctuation, number of resin ducts per tree-ring and position of resin ducts within the tree-rings. All parameters showed calendar-year patterns, visible as lines parallel to the bark. The most clear calendar-year pattern was seen for the type of transition from early to latewood and for intra-annual density fluctuations. The strongest inter-series correlation between calendar rings was seen for ring-width. None of the parameters showed significant inter-series correlations for cambial rings. These results may help us to understand how cores or discs taken at breast height represent the entire tree.     

Variation of different tree-ring parameters in samples from each terminal shoot of a Norway spruce tree

Variability of wood parameters in a tree is sometimes a rather nebulous concept since variability is evident within single cells, from early to latewood, from pith to bark and from stem base to the top of a tree. So far, stem analyses have been done using a restricted number of parameters, mostly ring-width, and using a restricted number of samples in the longitudinal direction. This study analyses a number of parameters from a single tree. An 81-year-old spruce tree was felled and internodial discs were taken from each annual terminal shoot. All tree rings in each disc were measured and a whole-stem analysis was completed for the following parameters: ring-width, mean ring density, maximum density, percentage of latewood, type of transition from early to latewood, intra-annual density fluctuation, number of resin ducts per tree-ring and position of resin ducts within the tree-rings. All parameters showed calendar-year patterns, visible as lines parallel to the bark. The most clear calendar-year pattern was seen for the type of transition from early to latewood and for intra-annual density fluctuations. The strongest inter-series correlation between calendar rings was seen for ring-width. None of the parameters showed significant inter-series correlations for cambial rings. These results may help us to understand how cores or discs taken at breast height represent the entire tree.     

Xylem water content and wood density in spruce and oak trees detected by high-resolution computed tomography

Elucidation of the mechanisms involved in long-distance water transport in trees requires knowledge of the water distribution within the sapwood and heartwood of the stem as well as of the earlywood and latewood of an annual ring. X-ray computed tomography is a powerful tool for measuring density distributions and water contents in the xylem with high spatial resolution. Ten- to 20-year-old spruce (Picea abies L. KARST.) and oak (Quercus robur) trees grown in the field were used throughout the experiments. Stem and branch discs were collected from different tree heights, immediately deep frozen, and used for the tomographic determinations of spatial water distributions. Results are presented for single-tree individuals, demonstrating heartwood and sapwood distribution throughout their entire length as well as the water relations in single annual rings of both types of wood. Tree rings of the sapwood show steep water gradients from latewood to earlywood, whereas those of the heartwood reflect water deficiency in both species. Although only the latest two annual rings of the ringporous species are generally assumed to transport water, we found similar amounts of water and no tyloses in all rings of the oak sapwood, which indicates that at least water storage is important in the whole sapwood.     

Relationship of strontium and calcium concentrations with the parameters of cell structure in Siberian spruce and fir tree-rings

This study addressed distribution of calcium and strontium in Siberian spruce (Picea obovata Ledeb.) and Siberian fir (Abies sibirica Ledeb.) tree-rings and its dependence on these woody species cell structure. Calcium concentration was found to decrease gradually from earlywood to latewood, whereas strontium showed an opposite trend. However, their trends at the scale of several rings are co-directed in the samples analyzed. A strong linear relationship was identified between the distribution of Sr/Ca concentration ratio and tree-ring density profile for both woody species. Radiographic density of Siberian spruce tree-ring cell walls and Ca and Sr concentrations in them were determined to have negative correlation with cell wall thickness. In earlywood of annual rings of a spruce the radiographic density of cell wall reaches 2.0 g/cm³ and decreases to 1.2 g/cm³ in latewood. The hypothesis put forward in this study to explain these strontium and calcium distributions in the tree-rings is that the concentrations of the element ions change with development of different cell wall layers. The high value of radiographic density of a cellular wall in earlywood and its relationship with cell wall thickness can be explained by the presence of ions of calcium in a cellular wall. Ions of calcium absorb X-ray radiation more strongly in comparison with light chemical elements. It can become the reason of observable relationship between radiographic density of cell wall and cell wall thickness.     

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.     

Effect of growth rate on mean microfibril angle and cross-sectional shape of tracheids of Norway spruce

The variation of the mean microfibril angle (MFA) and the shape of the cross-section of lumen with the distance from the pith in fast grown Norway spruce were studied by X-ray scattering and optical microscopy. The samples were from stems of a clone of Norway spruce [ Picea abies (L.) Karst.] grown in a fertile site at Nurmijärvi, southern Finland Both the mean MFA and the circularity index of the lumen of the fast-grown trees decreased more gradually as the distance from the pith increased than those in reference trees grown in a medium fertility site. However, in mature wood the mean MFA reached the same level in fast-grown trees as in reference trees (5°–10°) but the cross-sections of the cells remained more circular in fast-grown trees than in reference trees. The dependence of the mean MFA on the distance from the pith was similar for earlywood and latewood, but the values of the mean MFA of latewood were systematically smaller than those of earlywood. Two different X-ray diffraction geometries were compared from the points of view of biology and data analysis.     

False ring formation in eastern hemlock branches: impacts of hemlock woolly adelgid and elongate hemlock scale

Herbivores can alter plant physiology through the induction of abnormal wood formation. Feeding by some insects induces the formation of false rings, a band of thick-walled latewood cells within the earlywood portion of the tree ring that reduces water transport. Hemlock woolly adelgid (Adelges tsugae Annand) and elongate hemlock scale (Fiorinia externa Ferris) are invasive insects that both feed on eastern hemlock [Tsuga canadensis (L.) Carrière]. Adelges tsugae has a greater effect on tree health than F. externa, but the mechanism underlying their differential effect is unknown. We explored the effects of these herbivores by assessing growth ring formation in branches of trees that had been experimentally infested for 4 yr with A. tsugae, F. externa, or neither insect. We measured false ring density, ring growth, and earlywood: latewood ratios in the two most recently deposited growth rings. Branches from A. tsugae-infested trees had 30% more false rings than branches from F. externa-infested trees and 50% more than branches from uninfested trees. In contrast, branches from F. externa-infested trees and control trees did not differ in false ring formation. Radial growth and earlywood: latewood ratios did not differ among treatments. Our results show that two invasive herbivores with piercing-sucking mouth parts have differing effects on false ring formation in eastern hemlock. These false rings may be the product of a systemic plant hypersensitive response to feeding by A. tsugae on hemlock stems. If false rings are responsible for or symptomatic of hemlock water stress, this may provide a potential explanation for the relatively large effect of A. tsugae infestations on tree health.     

气候变暖背景下杉木年轮密度对气候因子的响应

为探讨杉木年轮密度与气候因子的响应关系,采用树木年轮学方法,以60年生杉木种源林为研究对象,测定杉木整轮密度、早材密度、晚材密度、晚材最大密度和早材最小密度,分析在气候变暖条件下主要气候因子（温度、降水、相对湿度）对杉木年轮密度及其生长的影响。结果表明：杉木不同年轮密度指标均受到温度、降水和相对湿度的显著影响。早材密度与当年夏季最高温度、当年5月降水量,最大密度与当年10月、当年秋季降水量,最小密度与前一年秋季降水量、最小相对湿度呈显著负相关。滑动相关分析表明气候因子在短时间尺度上对杉木生长影响的稳定性有显著影响,其中杉木年轮最大密度与当年10月、秋季的平均相对湿度和最小相对湿度,最小密度与当年2、3月的平均相对湿度和前一年秋季的平均相对湿度、最小相对湿度、降水量的负相关关系最为稳定。杉木年轮最小密度对前一年气候要素的响应存在滞后效应,且晚材密度对当年春季的气候要素响应也存在滞后效应。研究结果对开展亚热带针叶树种年轮生态学和年轮气候学研究具有重要参考价值,建议选择武夷山的天然林获取更长年表用于重建古气候。     

Annual cyclicity in high-resolution stable carbon and oxygen isotope ratios in the wood of the mangrove tree Rhizophora mucronata

In the present study, the high‐resolution stable carbon (¹³C/¹²C) and oxygen (¹⁸O/¹⁶O) isotope ratio profiles in the wood of the mangrove Rhizophora mucronata Lam., a tropical tree species lacking distinct growth rings, were investigated. Variations of both isotope ratios revealed a remarkable annual cyclicity with lowest values occurring at the latewood/earlywood boundary (April–May) and highest values during the transition from earlywood to latewood (October–November). Based on the current knowledge of the physiology of this mangrove species, as well as on the current literature available on high‐resolution profiles of stable isotope ratios in tree rings, possible driving forces responsible for this seasonal pattern are discussed. The annual cyclicity, together with a conspicuous isotope pattern appearing in the El‐Niño year 1997, promises great potential for tropical dendrochronology.     

Couplings in cell differentiation kinetics mitigate air temperature influence on conifer wood anatomy

Conifer trees possess a typical anatomical tree‐ring structure characterized by a transition from large and thin‐walled earlywood tracheids to narrow and thick‐walled latewood tracheids. However, little is known on how this characteristic structure is maintained across contrasting environmental conditions, due to its crucial role to ensure sap ascent and mechanical support. In this study, we monitored weekly wood cell formation for up to 7 years in two temperate conifer species (i.e., Picea abies (L.) Karst and Larix decidua Mill.) across an 8°C thermal gradient from 800 to 2,200 m a.s.l. in central Europe to investigate the impact of air temperature on rate and duration of wood cell formation. Results indicated that towards colder sites, forming tracheids compensate a decreased rate of differentiation (cell enlarging and wall thickening) by an extended duration, except for the last cells of the latewood in the wall‐thickening phase. This compensation allows conifer trees to mitigate the influence of air temperature on the final tree‐ring structure, with important implications for the functioning and resilience of the xylem to varying environmental conditions. The disappearing compensation in the thickening latewood cells might also explain the higher climatic sensitivity usually found in maximum latewood density.     

13CO2 pulse-labelling of photoassimilates reveals carbon allocation within and between tree rings

Post-photosynthetic fractionation processes during translocation, storage and remobilization of photoassimilate are closely related to intra-annual sigma13C of tree rings, and understanding how these processes affect tree-ring sigma13C is therefore indispensable for improving the quality of climate reconstruction. Our first objective was to study the relationship between translocation path and phloem grain. We pulse-labelled a branch of Larix gmelinii (Rupr.) Rupr. and later analysed the sigma13C distribution in the stem. A 13C spiral translocation path closely related to the spiral grain was observed. Our second objective was to study the use of remobilized storage material for earlywood formation in spring, which is a suspected cause of the autocorrelation (correlation of ring parameters to the climate in the previous year) observed in (isotope) dendroclimatology. We pulse-labelled whole trees to study how spring, summer and autumn photoassimilate is later used for both earlywood and latewood formation. Analysis of intra-annual sigma13C of the tree rings formed after the labelling revealed that earlywood contained photoassimilate from the previous summer and autumn as well as from the current spring. Latewood was mainly composed of photoassimilate from the current year's summer/autumn, although it also relied on stored material in some cases. These results emphasize the need for separating earlywood and latewood for climate reconstruction work with narrow boreal tree rings.     

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.     

Seasonal and perennial changes in the distribution of water in the sapwood of conifers in a sub-frigid zone

An analysis was made of progressive changes in patterns of cavitation in the sapwood of three species of conifer (Larix kaempferi, Abies sachalinensis, and Picea jezoensis) that were growing in a sub-frigid zone. In all three conifers, all tracheids of the newly forming outermost annual ring were filled with water or cytoplasm during the period from May to August. However, many tracheids in the transition zone from earlywood to latewood lost water in September, presumably through drought-induced cavitation. Cavitated tracheids tended to be continuously distributed in a tangential direction. Subsequently, some earlywood tracheids of the outermost annual ring lost water during the period from January to March. This was associated with freeze-thaw cycles. In the second and third annual rings from the cambium of all three conifers, the lumina of most tracheids in the transition zone from earlywood to latewood contained no water. In contrast, some latewood tracheids near the annual ring boundary and many earlywood tracheids retained water in their lumina. The third annual ring had more cavitated tracheids than the second annual ring. Our observations indicated that cavitation progressed gradually in the tracheids of the conifers and that they were never refilled once cavitation had occurred. The region involved in water transport in conifers did not include the entire sapwood and differed among annual rings.     

Effect of local heating and cooling on cambial activity and cell differentiation in the stem of Norway spruce (Picea abies)

BACKGROUND AND AIMS The effect of heating and cooling on cambial activity and cell differentiation in part of the stem of Norway spruce (Picea abies) was investigated. METHODS: A heating experiment (23-25 degrees C) was carried out in spring, before normal reactivation of the cambium, and cooling (9-11 degrees C) at the height of cambial activity in summer. The cambium, xylem and phloem were investigated by means of light- and transmission electron microscopy and UV-microspectrophotometry in tissues sampled from living trees. KEY RESULTS: Localized heating for 10 d initiated cambial divisions on the phloem side and after 20 d also on the xylem side. In a control tree, regular cambial activity started after 30 d. In the heat-treated sample, up to 15 earlywood cells undergoing differentiation were found to be present. The response of the cambium to stem cooling was less pronounced, and no anatomical differences were detected between the control and cool-treated samples after 10 or 20 d. After 30 d, latewood started to form in the sample exposed to cooling. In addition, almost no radially expanding tracheids were observed and the cambium consisted of only five layers of cells. Low temperatures reduced cambial activity, as indicated by the decreased proportion of latewood. On the phloem side, no alterations were observed among cool-treated and non-treated samples. CONCLUSIONS: Heating and cooling can influence cambial activity and cell differentiation in Norway spruce. However, at the ultrastructural and topochemical levels, no changes were observed in the pattern of secondary cell-wall formation and lignification or in lignin structure, respectively.     

Variation in wood anatomical structure of Douglas-fir defoliated by the western spruce budworm: a case study in the coastal-transitional zone of British Columbia,Canada

Key message

An outbreak of the western spruce budworm temporarily modifies cellular wood anatomy of stem wood in natural and mature Douglas-fir stands impacting wood quality properties.

Abstract

Western spruce budworm (Choristoneura occidentalis Freeman) is a widespread and destructive defoliator of commercially important coniferous forests in western North America. In British Columbia, Canada, Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] is the most important and widely distributed host. Permanent sample plots were established at a number of locations in southern interior at the beginning of a severe western spruce budworm outbreak in the 1970s. Two of the sites were sampled in 2012 to determine whether modifications had occurred in the anatomical characteristics of stem wood formed during outbreak years. We determined that rings formed during the western spruce budworm 1976–1980 outbreak had a significantly lower proportion of latewood, reduced mean cell wall thickness and smaller radial cell diameters. While the cellular characteristics of the earlywood remained fairly constant, significant reductions in lumen area occurred in 1978 and 1979 at each site. Our study shows that western spruce budworm outbreaks not only reduce annual radial growth, but also temporarily modify cellular characteristics in latewood cells, which has implications for wood density and quality in Douglas-fir.     

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.     

Hydraulic efficiency compromises compression strength perpendicular to the grain in Norway spruce trunkwood

The aim of this study was to investigate bending stiffness and compression strength perpendicular to the grain of Norway spruce (Picea abies (L.) Karst.) trunkwood with different anatomical and hydraulic properties. Hydraulically less safe mature sapwood had bigger hydraulic lumen diameters and higher specific hydraulic conductivities than hydraulically safer juvenile wood. Bending stiffness (MOE) was higher, whereas radial compression strength lower in mature than in juvenile wood. A density-based tradeoff between MOE and hydraulic efficiency was apparent in mature wood only. Across cambial age, bending stiffness did not compromise hydraulic efficiency due to variation in latewood percent and because of the structural demands of the tree top (e.g. high flexibility). Radial compression strength compromised, however, hydraulic efficiency because it was extremely dependent on the characteristics of the “weakest” wood part, the highly conductive earlywood. An increase in conduit wall reinforcement of earlywood tracheids would be too costly for the tree. Increasing radial compression strength by modification of microfibril angles or ray cell number could result in a decrease of MOE, which would negatively affect the trunk’s capability to support the crown. We propose that radial compression strength could be an easily assessable and highly predictive parameter for the resistance against implosion or vulnerability to cavitation across conifer species, which should be topic of further studies.     

Testing different Earlywood/Latewood delimitations for the establishment of Blue Intensity data: A case study based on Alpine Picea abies samples

For dendroclimatological Blue Intensity (BI) studies based on earlywood (EW) or latewood (LW) information, a demarcation between the two is necessary, which can be difficult to establish for species where the transition is subtle. Often, a percental value k is used that calculates an EW/LW boundary value for each tree ring individually based on the difference between maximum and minimum absorption. Several laboratories and authors have used different values for k (e.g. k = 30 % or k = 50 %), while wood anatomical and visual studies suggest that k is on the order of 80 %. Here, we test how different settings of k, and thus different definitions of the EW and LW proportions of a tree ring, influence the dendroclimatic potential of derived time series. To this end, we correlate instrumental temperature measurements with tree ring chronologies that are based on EW and LW information (e.g. EW absorption (EWBI), LW absorption (LWBI)), where the EW/LW proportion is varied by setting different values for k. The tree ring samples utilized are 30 cores of spruce (Picea abies) trees from a high-elevated site (ca. 1700 m a.s.l.) in the northern Alps, Austria. Overall, we achieve high correlations between temperature data and our tree ring chronologies. Regarding the stability of the climate signal under different k values, the results show that absorption intensity based parameters (ΔBI, EWBI, LWBI) are only mildly influenced by different settings of k, while width based parameters (EW width, LW width) show a larger dependence on k. LW width, for instance, was stronger correlated with temperature, the smaller the LW was chosen (and thus the higher k was set). Based on our results and the wood anatomical definition of the EW/LW boundary, we suggest that k = 80 % may be a good choice for future studies. However, since this is only a case study from one site, careful screening of the respective data set regarding an appropriate k value must accompany each dendroclimatological study.     

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。     

A comparison of two techniques for measuring and crossdating tree rings

Crossdating is the core principle of dendrochronology. Our study compared two techniques for measuring and crossdating tree rings using Juniperus virginiana L. (eastern redcedar) as a case study. We used a pseudo 2 × 2 study design comparing the traditional skeleton plot/sliding measuring stage technique to a semi-automatic image analysis program across two technicians. Crossdating was evaluated in COFECHA. Raw measurements of total, earlywood, and latewood widths from the two methods were analyzed using the Verify for Windows program, ANOVA, and correlation matrices. Total ring width and earlywood width were well correlated between techniques and technicians but questionable ring boundaries from image analysis program should be checked under a stereoscope. Juniperus virginiana latewood widths were significantly different between techniques and technicians; therefore, we do not recommend combining latewood measurements from species with limited latewood variability for dendrochronological analysis. A standard definition of the earlywood-latewood boundary that can be replicated across technicians is needed to combine latewood measurements from the sliding measuring stage and image analysis systems.