Duration and extension of anatomical changes in wood structure after cambial injury

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 excelsior L.) 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.     

Ensuring a representative sample of earlywood vessels for dendroecological studies: an example from two ring-porous species

The analysis of time series of wood cell anatomical features (such as the earlywood vessels of ring-porous trees) is a successful approach to understand the effect of environmental factors on tree growth and thus constitutes a valuable source of information about past environmental conditions. However, despite the rising interest in analyzing wood anatomical time series, little or no attention has been paid to establish an adequate sample of cells in order to minimize the risk of missing a valuable environmental signal. In order to contribute to such methodological bases, this paper is aimed at (1) identifying a representative sample of earlywood vessels within a tree, which encode the same climatic information, and (2) assessing if it is preferable to obtain the sample of vessels along one or two radii. Four individuals of sessile oak (Quercus petraea (Mattuschka) Liebl.) and sweet chestnut (Castanea sativa Mill.) were harvested and all their earlywood vessel lumina were measured along two 40-mm wide radial strips. Measured vessels were selected stepwise while increasing the tangential width of the wood section from 1 to 40 mm, analyzing at each step (1) the common signal of chronologies and (2) the correlation to the main climatic variables controlling growth. Additionally, both radii in each tree were analyzed together and separately. The results showed that a total tangential width of 10 mm was enough to stabilize the climatic signal with improvement when distributed along two different radii, but a slightly larger tangential width was required to reach an optimal common signal. We suggest that, at least for the case of these two species growing at this specific climatic context, two 5-mm increment cores ensure a representative vessel selection.     

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.     

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.     

Earlywood vessel size of oak as a potential proxy for spring precipitation in mesic sites

Aim In this study, we evaluate the importance of the mean earlywood vessel size of oaks as a potential proxy for climate in mesic areas. Location The study was conducted in Switzerland at three forest sites dominated by oak (Quercus petraea and Q. pubescens). The three sites were in different climatic zones, varying mainly in terms of precipitation regime. Methods Three 50‐year‐long site chronologies of mean earlywood vessel size and tree‐ring widths were obtained at each site and related to monthly meteorological records in order to identify the main variables controlling growth. The responses of mean vessel size to climate were compared with those of the width variables to evaluate the potential climatic information recorded by the earlywood vessels. Results The results show that the mean vessel size has a different and stronger response to climate than ring‐width variables, although its common signal and year‐to‐year variability are lower. This response is better in particular at mesic sites, where it is linked to precipitation during spring, i.e. at the time of vessel formation, and is probably related to the occurrence of only a few processes controlling vessel growth, whereas radial increment is controlled by multiple and varying factors. Main conclusions The mean earlywood vessel size of oak appears to be a promising proxy for future climate reconstructions of mesic sites, where radial growth is not controlled by a single limiting factor.     

An evaluation of english oak earlywood vessel area as a climate proxy in the UK

This research evaluates the usefulness of oak earlywood vessel area as a climate proxy in the western United Kingdom (UK). The results demonstrate that at this site earlywood vessel area contains a different environmental signal (March relative humidity) to a ring-width chronology developed from the same trees. The vessel area signal passes commonly used verification statistics and was found to be representative of the climate of a similar geographic area to other tree-ring proxies, albeit with a lower explained variance. Taking the average of all the vessels identified for each year weakened the reconstructed climate signal and it was found for this study that the average of the 10 largest vessels provided the strongest and most stable match. The results demonstrate earlywood vessel area of oak in the UK has potential as a climate proxy, but that further work to strengthen and characterise the climatic target variable controlling vessel area is required.     

Wood anatomy and tree growth covary in riparian ash forests along climatic and ecological gradients

Riparian ash forests subjected to seasonal drought are among the most endangered ecosystems in Europe. They are threatened by climate warming causing aridification and by land-use changes modifying river flow. To assess the impacts of these two stress factors on riparian forests, we studied radial growth and xylem anatomical traits in five narrow-leaved ash (Fraxinus angustifolia) stands across wide climatic and ecological gradients from northern Italy to southern Portugal. Radial growth rates and earlywood hydraulic diameter (Dh) were directly correlated, whilst earlywood vessel density and growth rates were inversely associated. Ash growth positively responded to precipitation. Higher and lower rates of growth increase in response to precipitation were found in dry (annual precipitation 357–750 mm, annual water balance −39 to −48 mm) and wet (annual precipitation 1030 mm, annual water balance 27 mm) sites, respectively. Wet conditions in autumn and winter of the year prior to tree-ring formation lead to larger Dh values, except in the wet site where warmer conditions from prior autumn to current spring were positively associated to wider vessels. Growth was also enhanced by a higher river flow, reflecting higher soil moisture due to elevated groundwater table levels. Peaks in river flow from late winter to early spring increased Dh in dry-continental sites. Growth and potential hydraulic conductivity in drought-prone riparian ash forests are differently impacted by climate variability and river flow depending on site and hydrological conditions. Nevertheless, covariation between radial growth and the earlywood vessel diameter was found, regardless of site specific differences. Wood production and hydraulic conductivity are coordinated through the production of large earlywood vessels which may allow reaching higher growth rates.     

Wood anatomy and vessel characteristics of spiny monkey orange (Strychnos spinosa) in Benin (West Africa)

The response of plant species to varying climate conditions in tropical Africa remains poorly understood but can be assessed using wood anatomical traits. These traits play an important role for the adaptive capacity of a species to environmental stress, since environmental conditions can modify the proportion, size, and morphology of wood anatomical elements. This study reports quantitative data on vessel characteristics of the diffuse porous angiosperm Spiny monkey orange (Strychnos spinosa Lam.) in Benin in tropical West Africa. The vessel-related anatomical traits varied with high amplitude (coefficient of variation CV ˃ 25%) between different sites located in different climate zones. The variability of the traits is higher within one climatic zone than between climatic zones, and even more pronounced within trees. Consequently, the climatic zones have less influence on the studied features than local site conditions. However, the study showed that S. spinosa individuals that have numerous vessels also have a high lumen fraction and total ring area. On the other hand, individuals presenting a high vessel density also display vessels of smaller size. The correlation between vessel number and total ring area on the one hand, and between vessel size and lumen fraction on the other hand are highly significant and positive. In Benin, S. spinosa wood anatomical traits are likely linked to local site factors rather than to regional climatic factors.     

Reactivation of Vessel Production in Ash (Fraxinus excelsior L.) Trees

Stages of earlywood vessel development have been compared withstages of bud and shoot growth throughout 12-year-old treesof ash (Fraxinus excelsior L.). Reactivation of vessel productionwas not simultaneous throughout the tree. There was evidencethat vessel expansion progressed basipetally down branches andacropetally up the main stem. The earliest expanding vesselswere found scattered around the circumference of main stem andbranches about 3 weeks before the emergence of foliage leavesfrom the buds. Other vessels expanded later between the earlierones so the whole of the first earlywood layer was expandingby 1.5 weeks before leaf emergence; this is suggested as a convenientstage to use as a baseline for a model of wood production inash. Vessel maturation progressed basipetally down the mainstem as well as branches, the first mature (presumably functional)vessels appearing in the upper stem shortly before leaf emergence.Mature vessels were not found in the lower part of the mainstem until after the beginning of rapid leaf expansion afterbudbreak, contrary to a previous assumption that functionalearlywood vessels are of necessity produced before leaf expansionin ring-porous trees. Patterns of vessel expansion are comparedbetween the ring-porous ash and the diffuse-porous sycamore;these data suggest that expansion of earlywood vessels beganat the same time in relation to budbreak in the two species,but the location of the first vessel expansion differed. ash, cambial reactivation, Fraxinus excelsior L., ring-porous hardwood, vessel expansion, vessel maturation     

Growth Rings,Phenology, Hurricane Disturbance and Climate in Cyrilla racemiflora L., a Rain Forest Tree of the Luquillo Mountains,Puerto Rico1

The growth phenology of Cyrilla racemiflora L., the dominant tree species of the montane rain forest, (subtropical lower montane rain forest, sensu Holdridge) of the Luquillo Mountains of Puerto Rico was studied intensively during 1989, and then semiannually through mid-1993 to determine the periodicity of changes in xylem structure. Four trees at 770 m were monitored for flowering, branch elongation, leaf litterfall, and xylem cell growth and differentiation in the lower stem, and these events were related to local seasonal patterns of rainfall and temperature. Hurricane Hugo defoliated study trees in September, 1989. Bud-break and branch elongation in March, 1989 were followed by earlywood xylem cell production in the lower stem in April and the onset of flowering in May. Leaf litterfall was greatest between April and June, coinciding with peak branch growth and new leaf formation. Latewood xylem was produced in December. The general phenological pattern was synchronized between trees and over study years. Vessel diameter and density were monitored along with thickness of earlywood and latewood and the former converted to vessel lumen area, a measure of xylem conductance capacity. Annual growth rings were formed with periods of earlywood and latewood production coinciding with traditional summer (rainy) and winter (dry) seasons, respectively, in the Luquillo Mountains. Hurricane defoliation was followed by heavy flowering in 1990, a year of reduced branch elongation and annual xylem ring width, and was associated with maximum vessel lumen area, as was flowering in 1989, prior to the hurricane. Hurricane Hugo provided a perturbation that, through its elicited stress response, allowed for the demonstration of the interplay between flowering, branching, structural growth of xylem, and xylem function.     

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.     

Xylem water-conducting patterns of 34 broadleaved evergreen trees in southern Japan

A dye injection method was used to elucidate the xylem water-conducting pathways of 34 broadleaved evergreen trees growing in southern Japan: two semi-ring-porous, 26 diffuse-porous, five radial-porous and one non-vessel species. The large earlywood vessels in semi-ring-porous species have a water transport function in only the outermost annual ring, as in deciduous ring-porous species. On the other hand, the small vessels in semi-ring-porous species maintain the water transport function in many outer annual rings. For the other xylem-type species, the many vessels in many outer annual rings have a water transport function. In diffuse-porous species, we categorized the water-conducting pattern within the annual rings into two types: d1 type, where water travels through vessels in the whole region; and d2 type, where water travels mainly through the earlywood vessels. The pattern in radial-porous species is similar to that in the d1 type; the pattern in non-vessels species is similar to that in the d2 type. The vessel diameter in radial-porous species is similar to that of the earlywood vessels of semi-ring-porous species. These results suggest that the conduit diameter size is only one of many factors determining the water-conducting pathways of broadleaved evergreen species.     

Wood anatomical changes in roots of European ash (Fraxinus excelsior L.) after exposure

Global changes have distinct influence on fluvial processes in torrents causing erosion on slopes and riverbanks even in forested areas. Continuous as well as discontinuous erosion of riverbanks covered by trees frequently results in destabilisation and finally cause the trees affected to tip over. These uprooted trees may lead to a blockage of the river and, in the case of a collapse of the resulting dam, resulting in severe flooding or even debris-flow surges. Dating the time of root exposure along riverbanks allows the reconstruction of erosion dynamics. In the previous studies, roots of coniferous trees have shown anatomical changes (cell size reduction in earlywood) after exposure. This anatomical feature in the xylem of exposed roots has helped to determine the time of exposure at an annual resolution. In the current study, this methodology is now applied for the first time to European ash (Fraxinus excelsior L.) to determine its specific reactions to root exposure.

First results show different reactions in vessel and fibre dimensions. Fibres show a distinct decrease in lumen area due to exposure, whereas vessel lumen area displays more variability ranging from 50% size reduction to no changes. It is discussed that vessel size characteristics are also related to ecophysiological and climatic patterns.     

FLUID FLOW IN THE OUTERMOST XYLEM INCREMENT OF A RING-POROUS TREE,ULMUS AMERICANA

Hydraulic conductivity through the outermost growth ring of Ulmus americana was quantified to determine the significance of this single growth increment to fluid flow in xylem of a ring-porous tree. Gravity flow rates through trunks deprived of the outermost growth ring dropped to 8% of the rate achieved in control trunks whose cross section was intact. However, colored dye, fed through stem segments, appeared in up to 4-yr old wood, corroborating earlier reports that fluid flows through more than just the outermost growth ring of ring-porous trees. Finally, the pathway of flow through older rings was shown to consist of narrow latewood elements while the wider and more hydraulically significant earlywood vessels came into play only in the outermost growth ring of U. americana. These data indicate that the outermost growth ring of Ulmus is responsible for over 90% of xylem transport in this ring-porous tree, because wide earlywood vessels function only for one growth season.     

Myxomycetes developed on litter of common Finnish trees in moist chamber cultures

Fallen leaves were collected from the Finnish trees Alnus gtutinosa & A. incana, Betula pendula & B. pubescens, Picea abies, Pinus sylvestris and Populus tremula , and used to establish 400 moist chamber cultures. From these emerged 132 specimens of Myxomycetes belonging to 14 species. One of them, Didymium dubium Rostaf. was also cultivated from spore to spore in artificial media. The three Arcyria species that emerged developed only on coniferous litter and the four Didymium species only on deciduous litter. Echinostelium minitum de Bary is the only myxomycete species that emerged on leaves of all the five trees.
Alnus litter gave rise to the greatest number of myxomycete specimens. Betula leaves seemed to be the most unfavourable of the substrates. The common species of Myxomycetes that have appeared only on bark of living trees in the moist chambers set up by the author include Paradiacheopsis fimbriata (G. Lister & Cran) Hertel and P. solitaria (Nann.–Brem.) Nann.–Brem. Species occurring only on deciduous leaves or grains are Didymium difforme (Pers.) S. F. Gray, D. dubium Rost. and D. squamulosum (Alb. & Schw.) Fr.     

不同林分类型林缘华北落叶松细胞特征

以塞罕坝机械林场华北落叶松纯林及华北落叶松-白桦混交林为对象,研究林地边缘华北落叶松边界木与林内木径向生长、年轮细胞大小及数量的差异,分析林缘对不同林分类型华北落叶松径向生长及细胞特征的影响。结果表明: 纯林中边界木的径向生长显著快于林内木,年轮总宽、早材和晚材总生长量在边界木较林内木分别增加48.9%、58.9%和29.6%,而混交林边界木与林内木的径向生长差距并不明显。纯林边界木早材细胞总数、早材大细胞和小细胞数量较林内木分别增加63.3%、55.6%和70.0%,晚材细胞总数、晚材大细胞和小细胞数量边界木较林内木分别增加35.4%、37.5%和28.5%,而早晚材细胞大小在边界木与林内木间无显著差异。混交林边界木早晚材细胞数量与林内木无显著差异,但边界木早材细胞大小较林内木增大50.0%,边界木早材最大细胞、最小细胞、大细胞及小细胞的大小较林内木分别增大28.6%、33.3%、16.6%和25.0%。通过混交的方式营造混交林可以有效地缓解林缘导致的纯林中边界木生长过快而林内木生长较慢的现象。     

Tree-ring structure and climatic effects in young Eucalyptus globulus Labill. grown at two Portuguese sites: preliminary results

Dendrochronological analyses of Eucalyptus are generally rare and mostly restricted to Australia. This work reports an attempt to identify annual ring patterns in young blue gum trees (Eucalyptus globulus Labil) grown in Europe through studying the radial variation of vessel characteristics. Common radial trends were seen such as increasing vessel areas, decreasing vessel abundances and a cyclic variability of vessel characteristics associated to the seasons. Low water availability was related to reduced ring widths in combination with more and smaller vessels. Even with the given juvenility of the investigated eucalypt trees the pith to bark trends of vessel parameters might be useful in dendroecological studies, particularly with respect to drought scenarios. The employed method of using vessel parameters to identify rings seem to be the only accurate way of identifying annual rings in E. globulus.     

Growth reactions of Pinus sylvestris L. and Quercus pubescens Willd. to drought years at a xeric site in Valais, Switzerland

In Valais, an inner-Alpine dry valley in Switzerland, low-elevation Scots pine (Pinus sylvestris L.) forests are changing. While pine shows high mortality rates, deciduous species, in particular pubescent oak (Quercus pubescens Willd.), are becoming more abundant. We hypothesise that increasing drought and the species-specific drought tolerance are key factors in these processes. In this study, the growth reaction to drought years of pine and oak growing at a xeric site in Valais was analysed using dendrochronological and wood anatomical methods. Congruent with theoretical expectations, the tree-ring widths of both species, the mean lumen area of earlywood vessels in oak and the number of tracheids in a radial row in pine decreased in response to dry conditions. However, both species also showed reactions deviating from those known from mesic sites: In oak, the mean lumen area of latewood vessels increased in drought years. In pine, in the driest year of the period (1976), the mean radial diameter increased in latewood and decreased only slightly in earlywood. These results emphasises that the process of wood formation and cell functionality at xeric sites is not completely understood yet. Both species seem to have difficulties to adapt the size of their water-conducting cells to strongly reduced water availability in drought years. Additionally, the cell number is strongly reduced. Thus it remains unclear if both species can maintain sufficient water transport under increasingly dry conditions.     

The occurrence of vascular tracheids inBetula and some otherBetulaceae genera

The occurrence of vascular tracheids inBetula, Alnus, Carpinus andCorylus is reported. It is established that the cells are a consistent feature in early and late wood where they are associated with vessel elements in radial multiples. The possible function and taxonomic importance of these cells are briefly considered.     

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.