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Aim Spatial variations of environmental conditions translate into biogeographical patterns of tree growth. This fact is used to identify the origin of timber by means of dendroprovenancing. Yet, dendroprovenancing attempts are commonly only based on ring‐width measurements, and largely neglect additional tree–ring variables. We explore the potential of using wood anatomy as a dendroprovenancing tool, and investigate whether it increases the precision of identifying the origin of oak wood. Since different tree–ring variables hold different information on environmental conditions prevailing at specific times of the growing season—which vary between source regions—we hypothesize that their inclusion allows more precise dendroprovenancing. Location Europe, Spain. Taxon Quercus robur L., Quercus petraea (Matt.) Liebl., Quercus faginea Lam., Quercus pyrenaica Willd. Methods We sampled four oak species across Northern Spain, i.e. from the Basque country and Cantabria and—in the Basque country—from low to high elevation (topographic/latitudinal gradient). We measured multiple tree–ring variables to (a) extract complementary variables; (b) present statistical relations among them; (c) analyse region‐specific variation in their patterns based on time–series of individual trees; and (d) determine underlying climate–growth relationships. Leave‐one‐out analysis was used to test whether a combination of selected variables allowed dendroprovenancing of a randomly selected tree within the area. Results A combination of latewood width (LW) and earlywood vessel size was used to pinpoint the origin of oak wood with higher precision than ring width or LW only. Variation in LW pinpointed the wood to east and west areas, whereas variation in vessels assigned wood to locations along a latitudinal/topographic gradient. The climatic triggers behind these gradients are respectively an east–west gradient in June–July temperature and a north–south gradient in winter/spring temperatures. The leave‐one‐out analyses supported the robustness of these results. Main conclusions Integration of multiple wood–xylem anatomical variables analysed with multivariate techniques leads to higher precision in the dendroprovenancing of ring‐porous oak species.

We monitored leaf phenology and xylogenesis of 12 Pedunculate oaks in northern Poland in 2014. We hypothesized that the individual trees, which differed in size, age and habitat (tree stand or gap), also diverged in terms of the seasonal patterns of leaf phenology and xylogenesis. The samples used for wood formation observations were collected most frequently during the early leaf phenophases (from March to end of June). The transverse sections of the cambial region were cut with a sledge microtome. We counted the number of cambial cells, measured the width of xylem increment and assessed the timing of xylogenesis and earlywood-vessel formation. We found significant differences in leaf phenology and timing of xylogenesis among individual trees. The smallest differences in wood formation among the trees were observed at the beginning of the vegetation season when the first earlywood vessels were detected (9 days). The dates of completion of the first tangential row of earlywood vessels varied by up to 30 days, while for the completion of the entire earlywood dates varied by up to 32 days. The highest productivity of cambial cells (13 cell layers) was observed around the time of bud swelling at mid-April. In the last days of April, the number of cambial cell layers decreased and subsequently increased again when the leaves were nearly fully expanded at the end of May. To summarize, we observed a high seasonal variability in the number of cambial cell layers. Differences in the time of cessation of cambial activity and xylogenesis amounted to 1 month. We conclude that: (1) oak tree-ring widths and earlywood-vessel sizes and numbers may not be sensitive indicators for early spring temperature and spring defoliation; (2) the missing association between leaf phenophases and xylogenesis as well as the phenological variability may be the reasons for the lack of a clear climatic effect on the above-mentioned parameters.

Key messageA gradual change exists in the altitudinal response of earlywood growth of Abies fargesii to hydrothermal conditions, with temperature being the main climatic factor controlling its latewood growth.The Tibetan Plateau, as the “Third Pole”, has witnessed profound and intricate effects of climate change in recent decades. This may result in different responses of tree radial growth to climatic factors in this region, varying with elevation and growth stages. To accurately reveal these different responses we established totalwood, earlywood and latewood width chronologies of Abies fargesii at four elevations. Our results showed that: (1) As the elevation increases, the response of radial growth of Abies fargesii to climatic factors shifted from restriction primarily by water deficiency caused by rapid warming, to weak restriction by winter precipitation with favorable hydrothermal conditions, and then to double restriction owing to insufficient heat and excessive moisture. This pattern was inconsistent with the relationship between the radial growth and climatic factors for latewood observed at the low and middle–low elevations. (2) There existed a temporal variability in the relationship between tree radial growth and restrictive climate factors. This unstable relationship was mainly observed in the middle–low elevation within suitable ecological conditions and during the later stages of tree radial growth with lower growth rates. This is related to the fact that more suitable ecological conditions and lower growth rates make tree growth more susceptible to climatic fluctuations. Considering these findings, if the climate would experience further cooling and humidity increasing in the study region, the growth of Abies fargesii might be more adversely affected at high elevation. For economic and ecological considerations, the afforestation of Abies fargesii should be prioritized in the middle–low elevation zones where favorable for the population distribution.

Cambial injury has been reported to alter wood structure in broad-leaved trees. However, the duration and extension of associated anatomical changes have rarely been analysed thoroughly. A total of 18 young European ash (Fraxinus excelsiorL.) trees injured on the stem by a spring flood were sampled with the aim of comparing earlywood vessels and rays formed prior to and after the scarring event. Anatomical and hydraulic parameters were measured in five successive rings over one-quarter of the stem circumference. The results demonstrate that mechanical damage induces a decrease in vessel lumen size (up to 77%) and an increase in vessel number (up to 475%) and ray number (up to 115%). The presence of more earlywood vessels and rays was observed over at least three years after stem scarring. By contrast, abnormally narrow earlywood vessels mainly developed in the first ring formed after the event, increasing the thickness-to-span ratio of vessels by 94% and reducing both xylem relative conductivity and the index for xylem vulnerability to cavitation by 54% and 32%, respectively. These vessels accumulated in radial groups in a 30° sector immediately adjacent to the wound, raising the vessel grouping index by 28%. The wound-induced anatomical changes in wood structure express the functional need of trees to improve xylem hydraulic safety and mechanical strength at the expense of water transport. Xylem hydraulic efficiency was restored in one year, while xylem mechanical reinforcement and resistance to cavitation and decay lasted over several years.

Interannual variability of wood density – an important plant functional trait and environmental proxy – in conifers is poorly understood. We therefore explored the anatomical basis of density. We hypothesized that earlywood density is determined by tracheid size and latewood density by wall dimensions, reflecting their different functional tasks. To determine general patterns of variability, density parameters from 27 species and 349 sites across the Northern Hemisphere were correlated to tree-ring width parameters and local climate. We performed the same analyses with density and width derived from anatomical data comprising two species and eight sites. The contributions of tracheid size and wall dimensions to density were disentangled with sensitivity analyses. Notably, correlations between density and width shifted from negative to positive moving from earlywood to latewood. Temperature responses of density varied intraseasonally in strength and sign. The sensitivity analyses revealed tracheid size as the main determinant of earlywood density, while wall dimensions become more influential for latewood density. Our novel approach of integrating detailed anatomical data with large-scale tree-ring data allowed us to contribute to an improved understanding of interannual variations of conifer growth and to illustrate how conifers balance investments in the competing xylem functions of hydraulics and mechanical support.

Key messageUnder drier spring conditions, Q. variabilis might slow the FEV growth rate but will not produce narrower FEVs. FEV size might be a trade-off between hydraulic efficiency and hydraulic safety.Wood anatomical features can encode tree reactions to fluctuating environmental conditions. The first row of earlywood vessels (FEV) has proven to be very promising with respect to the inter-annual climate variability. However, knowledge of how intra-annual FEV formation reflects climate is still limited. We observed wood formation of Quercus variabilis in a warm temperate–subtropical climate transition at Qinling Mountain, China. Using micro-cores, growth of six trees was monitored at weekly/biweekly intervals February–December in 2015 and 2016, of which 2016 had a drier spring. Cambial activity and xylem cell differentiation were documented by microscopic observation. The FEV diameter and ring width were measured. The onset of xylem growth, which significantly differed between the years, occurred 8 days earlier in 2016, in mid-March. However, the completion of FEV was synchronized in mid-April over the 2 years. This delayed completion of FEV in 2016, when precipitation increased and Standardized Precipitation Evapotranspiration Index changed from a negative to positive value. The cessation of xylem formation was 3 weeks earlier during the drought year (September 5, 2016) compared to the moist year (September 29, 2015) because of the drought conditions in late summer. This finding was supported by evidence that a positive and significant correlation between precipitation and xylem growth was found only in 2016. The ring width differed significantly between the years, but the FEV diameter did not. Our results suggest that the drought-induced delayed formation of FEV and earlier cessation could be a strategy for oak trees to adapt to water stress.

Conifer tree rings are generally composed of large, thin-walled cells of light earlywood followed by narrow, thick-walled cells of dense latewood. Yet, how wood formation processes and the associated kinetics create this typical pattern remains poorly understood. We monitored tree-ring formation weekly over 3 yr in 45 trees of three conifer species in France. Data were used to model cell development kinetics, and to attribute the relative importance of the duration and rate of cell enlargement and cell wall deposition on tree-ring structure. Cell enlargement duration contributed to 75% of changes in cell diameter along the tree rings. Remarkably, the amount of wall material per cell was quite constant along the rings. Consequently, and in contrast with widespread belief, changes in cell wall thickness were not principally attributed to the duration and rate of wall deposition (33%), but rather to the changes in cell size (67%). Cell enlargement duration, as the main driver of cell size and wall thickness, contributed to 56% of wood density variation along the rings. This mechanistic framework now forms the basis for unraveling how environmental stresses trigger deviations (e. g. false rings) from the normal tree-ring structure

Narrow-leaved ash (Fraxinus angustifolia Vahl), a highly valued European forest tree species, has been severely affected by a large-scale decline, which is most probably driven by a complex of multiple interacting factors including fungi, which contribute to and accelerate this process. Red deer (Cervus elaphus L.) can be considered as one of the contributing factors, as they inflict damage on the stems of young trees by stripping the bark. These wounds not only represent suitable entry points for fungi which can cause tissue necroses and decomposition, they can lead to changes in the wood structure as well. The aims of this research were to analyze chosen parameters of bark stripping wounds in narrow-leaved ash stands, identify fungi present in the tissue exposed by wounding, and inspect the effect of wounding on the wood structure. Bark stripping was observed on ash trees from 2 cm up to 18 cm of DBH and between 0.1 m and 1.9 m of stem height. The most susceptible trees were those with an average DBH of 5 ± 2.5 cm. On most of the ash trees (51%), one-third to two-thirds of the circumference was damaged. In wounded tissue, 174 fungal isolates were found, most of which belonged to known endophytic fungi from the genera Trichoderma, Fusarium, and Clonostachys. It was observed that earlywood cells in the wounding zone had narrower lumens compared to ones in adjacent healthy zone with regard to different trees and stem heights.

Flowering plants predominantly conduct water in tubes known as vessels, with vessel diameter playing a crucial role in plant adaptation to climate and reactions to climate change. The importance of vessels makes it essential to understand how and why vessel diameter, plant height, and other ecological factors are interrelated. Although shoot length is by far the main driver of variation in mean vessel diameter across angiosperms, much remains to be understood regarding the factors accounting for the abundant variation around the y-axis in plots of mean species vessel diameter against shoot length. Here, we explore the potential role of porosity types, wood density, leaf phenology, background imperforate tracheary element type, vasicentric tracheids, vascular tracheids, perforation plate type, and successive cambia in causing variation in the y-intercept or slope of the mean species vessel-diameter– and vessel-density–shoot-length associations at the shoot tip and base. We detected numerous cases of ecologically significant variation. For example, latewood vessels of ring porous species scale with a lower slope than earlywood, i.e., latewood vessels are relatively narrow in taller plants. This pattern is likely the result of selection favoring freezing-induced embolism resistance via narrow vessels. Wood density was negatively associated with vessel diameter, with low wood density plants having wider vessels for a given height. Species with scalariform perforation plates scale with a lower shoot base vessel-diameter–shoot-length slope, likely reflecting selection against scalariform plates in wide vessels. In other cases, functional groups scaled similarly. For example, species with successive cambia did not differ from those with conventional single cambia in their mean vessel-diameter–shoot-length scaling, rejecting our prediction that species with successive cambia should have narrower vessels for a given shoot length. They did, however, have fewer vessels per unit shoot cross-sectional area than plants of similar heights, likely because vessels have longer functional lifespans (and therefore are fewer) in species with successive cambia. Our methods illustrate how vessel diameter can be studied taking shoot length into account to detect ecologically important variation and construct theory regarding plant adaptation via the hydraulic system that includes plant size as a vital element.

Many researches have been conducted to investigate creep behavior of wood; however, the effects of structure on wood creep behavior remain unclear. Therefore, the effects of existence and distribution of earlywood vessel belt on creep behavior of white oak ( Quercus alba L.) wood were investigated by dynamic thermal mechanical analyzer (DMA) with double cantilever bending in this study. Besides, a comparative numerical modeling simulation on strain curves of white oak specimens was completed using Burger and Five-parameter model. Results revealed that instantaneous strain and 45-min strain of specimens decreased with increase in the distance between earlywood vessel belt and stress acting surface obviously. Additionally, instantaneous strain and 45-min strain of specimens remarkably increased with increase in temperature from 20 to 80 °C. An obvious bending creep behavior was observed with increase in temperature from 20 to 80 °C. Both Burger and Five-parameter model can effectively simulate the creep behavior of white oak specimens with R 2 values greater than 0.90. Furthermore, Five-parameter model illustrated a better fitting effect than Burger model in the final creep stage due to the introduction of a non-linear creep strain growth expression. It concluded that creep behavior of white oak wood strongly depends on the existence and distribution of earlywood vessel belt.