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.     

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.     

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.     

Defining an adequate sample of earlywood vessels for retrospective injury detection in diffuse-porous species

Vessels of broad-leaved trees have been analyzed to study how trees deal with various environmental factors. Cambial injury, in particular, has been reported to induce the formation of narrower conduits. Yet, little or no effort has been devoted to the elaboration of vessel sampling strategies for retrospective injury detection based on vessel lumen size reduction. To fill this methodological gap, four wounded individuals each of grey alder (Alnus incana (L.) Moench) and downy birch (Betula pubescens Ehrh.) were harvested in an avalanche path. Earlywood vessel lumina were measured and compared for each tree between the injury ring built during the growing season following wounding and the control ring laid down the previous year. Measurements were performed along a 10 mm wide radial strip, located directly next to the injury. Specifically, this study aimed at (i) investigating the intra-annual duration and local extension of vessel narrowing close to the wound margin and (ii) identifying an adequate sample of earlywood vessels (number and intra-ring location of cells) attesting to cambial injury. Based on the results of this study, we recommend analyzing at least 30 vessels in each ring. Within the 10 mm wide segment of the injury ring, wound-induced reduction in vessel lumen size did not fade with increasing radial and tangential distances, but we nevertheless advise favoring early earlywood vessels located closest to the injury. These findings, derived from two species widespread across subarctic, mountainous, and temperate regions, will assist retrospective injury detection in Alnus, Betula, and other diffuse-porous species as well as future related research on hydraulic implications after wounding.     

Expeditious building of ring-porous earlywood vessel chronologies without loosing signal information

Chronologies of earlywood vessel size of ring-porous trees contain valuable ecological information, but long preparation procedures limit their application in ecological studies. Recent and fast techniques for wood surface preparation combined with automated image analysis are reducing the work needed to build chronologies, but might also entail measurement inaccuracy. In this study, we aim to evaluate the effect of a possible efficiency-accuracy trade-off on ecological signal strength. To this end, we compare measurements of mean vessel area from two recent and fast procedures carried out on sanded wood surfaces with a reference procedure based on an accurate survey from thin sections. Measurements were performed on increment cores of 15 sessile oaks (Quercus petraea (Mattuschka) Liebl.) for the period 1956–2006. Dissimilarities in results with the reference procedure were quantified and evaluated. Our data show that the workload can be reduced by more than 20-fold when using the highly automated procedure. Signal weakening caused by measurement errors is negligible for vessels >6,000 μm² and can be easily compensated by increasing the sample size. Manual correction of misrecognized vessels hardly reduced this error further. The new procedures constitute a major step towards an efficient and accurate analysis of earlywood vessel chronologies of ring-porous tree species.     

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.     

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.     

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     

Investigation of the influence of blood flow rate on large vessel cooling in hepatic radiofrequency ablation.

Radiofrequency (RF) ablation using high-frequency current has become an important treatment method for patients with non-resectable liver tumors. Tumor recurrence is associated with tissue cooling in the proximity of large blood vessels. This study investigated the influence of blood flow rate on tissue temperature and lesion size during monopolar RF ablation at a distance of 10 mm from single 4- and 6-mm vessels using two different approaches: 1) an ex vivo blood perfusion circuit including an artificial vessel inserted into porcine liver tissue was developed; and 2) a finite element method (FEM) model was created using a novel simplified modeling technique for large blood vessels. Blood temperatures at the inflow/outflow of the vessel and tissue temperatures at 10 and 20 mm from the electrode tip were measured in the ex vivo set-up. Tissue temperature, blood temperature and lesion size were analyzed under physiological, increased and reduced blood-flow conditions. The results show that changes in blood flow rate in large vessels do not significantly affect tissue temperature and lesion size far away from the vessel. Monopolar ablation could not produce lesions surrounding the vessel due to the strong heat-sink effect. Simulated tissue temperatures correlated well with ex vivo measurements, supporting the FEM model.     

Earlywood vessels of the sub-Mediterranean oak Quercus pyrenaica have greater plasticity and sensitivity than those of the temperate Q. petraea at the Atlantic–Mediterranean boundary

Key message

Earlywood vessel features indicate different adaptations of Quercus petraea and Q. pyrenaica , which are probably related with their corresponding Atlantic and sub-Mediterranean ecological requirements.

Abstract

We studied the climatic signal of the earlywood anatomy of a temperate [Quercus petraea (Mattuschka) Liebl.] and a sub-Mediterranean (Quercus pyrenaica Willd.) oak species growing under similar climatic conditions in a transitional area between the Atlantic and Mediterranean regions of the Iberian Peninsula. We hypothesized that both species react differently in their wood anatomy due to their contrasting ecological requirements, and we test the usefulness of earlywood anatomical features to study the behaviour of these ring-porous oaks upon climate. For this, we measured the earlywood vessels, and obtained annual series of several anatomical variables for the period 1937–2006 using dendrochronological techniques, considering whether the vessels belonged to the first row or not. After optimizing the data set by principal component analysis and progressive filtering of large vessels, we selected maximum vessel area and total number of vessels as they resulted to be the optimal variables to describe vessel size and number, respectively. Vessel size of Q. pyrenaica was dependent on precipitation along the previous growing season, whereas it did not show any clear climatic response for Q. petraea. On the contrary, vessel number was related to winter temperature for both species. These relationships observed between climate and anatomy appeared to be stable through time. The results obtained reinforce the utility of earlywood vessel features as potential climate proxies.     

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.     

水热耦合对沈阳地区油松木质部生长的影响

利用微树芯技术可以从细胞尺度研究树木形成层物候和径向生长的过程,揭示树木生长与气候的关系。油松是我国北方森林的建群树种之一,也是沈阳地区的优势树种。研究2020年整个生长季(4—11月)油松周尺度的形成层及木质部细胞变化,分析油松在沈阳地区的生长规律。结果表明: 油松形成层分裂活动开始于4月初,结束于9月末。木质部从扩大细胞出现(4月中旬)开始生长,到木质化细胞消失(10月下旬)结束,其生长符合“S”型曲线。2020年生长53个/列木质部细胞,最大生长速率(0.55个/列/d)出现在5月末,早晚材细胞于7月末发生转变。在沈阳地区最低温达到0 ℃以上时树木形成层开始活动,影响木质部生长开始和结束的最低临界温度为2～3 ℃。降水在油松整个生长过程中起到促进作用。沈阳地区7月末的高温和水分供给不足是木质部细胞分化形成早晚材的主要因子。     

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.     

The Progression of Cavitation in Earlywood Vessels of Fraxinus mandshurica var japonica during Freezing and Thawing

For an examination of the progression of cavitation in large-diameter earlywood vessels of a deciduous ring-porous tree, potted saplings of Fraxinus mandshurica var japonica Maxim. were frozen and then thawed. The changes in the amount and distribution of water in the lumina of the current year's earlywood vessels during the course of the freezing and thawing were visualized by cryo-scanning electron microscopy. When samples were frozen, most of the current year's earlywood vessels were filled with water. After the subsequent thawing, the percentage of cavitated current-year earlywood vessels gradually increased with time. All of the current year's earlywood vessels were cavitated within 24 h, and only limited amounts of water remained in the lumina of earlywood vessels. Similar cavitation of earlywood vessels was observed after thawing of frozen, excised stem pieces. In contrast, many vessels of the current year's latewood retained water in the lumina during freezing and thawing. These observations indicate that the cavitation of the current year's earlywood vessels is not produced during freezing but progresses during rewarming after freezing in F. mandshurica var japonica.     

The effects of localized heating and disbudding on cambial reactivation and formation of earlywood vessels in seedlings of the deciduous ring-porous hardwood,Quercus serrata

Background and Aims

The networks of vessel elements play a vital role in the transport of water from roots to leaves, and the continuous formation of earlywood vessels is crucial for the growth of ring-porous hardwoods. The differentiation of earlywood vessels is controlled by external and internal factors. The present study was designed to identify the limiting factors in the induction of cambial reactivation and the differentiation of earlywood vessels, using localized heating and disbudding of dormant stems of seedlings of a deciduous ring-porous hardwood, Quercus serrata.

Methods

Localized heating was achieved by wrapping an electric heating ribbon around stems. Disbudding involved removal of all buds. Three treatments were initiated on 1 February 2012, namely heating, disbudding and a combination of heating and disbudding, with untreated dormant stems as controls. Cambial reactivation and differentiation of vessel elements were monitored by light and polarized-light microscopy, and the growth of buds was followed.

Key Results

Cambial reactivation and differentiation of vessel elements occurred sooner in heated seedlings than in non-heated seedlings before bud break. The combination of heating and disbudding of seedlings also resulted in earlier cambial reactivation and differentiation of first vessel elements than in non-heated seedlings. A few narrow vessel elements were formed during heating after disbudding, while many large earlywood vessel elements were formed in heated seedlings with buds.

Conclusions

The results suggested that, in seedlings of the deciduous ring-porous hardwood Quercus serrata, elevated temperature was a direct trigger for cambial reactivation and differentiation of first vessel elements. Bud growth was not essential for cambial reactivation and differentiation of first vessel elements, but might be important for the continuous formation of wide vessel elements.     

Climatic control of intra-annual wood density fluctuations of Pinus pinaster in NW Spain

Tree rings of Pinus pinaster often contain intra-annual density fluctuations (IADFs), which have been attributed to the succession of dry and rainy periods typical of Mediterranean climate, but their formation has not been studied yet under Atlantic climate. We analyzed the occurrence and climatic significance of replicated IADFs in ten monospecific stands in NW Spain. The frequency of IADFs was higher than previously reported for this species under Mediterranean conditions and consistently decreased with increasing elevation. The formation of bands of latewood-like tracheids within the earlywood was favored by dry previous August, cold previous winter and dry April. Bands of earlywood-like tracheids within the early latewood were also favored by low winter temperatures. However, their occurrence was geographically heterogeneous, with two groups of stands being defined by their distances to the shoreline. In coastal stands, cold May–August triggered IADFs formation, while in inland stands their formation was favored by dry May–July. Regional winter temperatures and April water balance were strongly related to the East Atlantic (EA) pattern, which greatly conditioned the occurrence of IADFs in the earlywood and the early latewood. By contrast, the presence of bands of earlywood-like tracheids in the late latewood was independent of the EA pattern, being strongly related to warm conditions in spring and especially to a wet October. The link between regional climate and the EA pattern strongly controlled the physiological processes that determine intra-annual growth dynamics and short-term cell enlargement of P. pinaster in NW Spain.     

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.     

Chronologies of earlywood vessels and latewood width disentangle climate drivers of oak growth in a mild oceanic region

We obtained tree-ring chronologies for pedunculate oak along an altitudinal gradient of four sites in NW Iberia, which covers most of the species range within the region. Trees grow under Atlantic climate, with a rainfall maximum in autumn-winter, and a minimum during summer, but lacking a remarkable drought. Chronologies included several earlywood anatomical features and ring widths, which were compared to each other. Latewood width and a subset of six earlywood variables were selected for further analyses, considering the period 1954–2003.The lowest site considerably differed from the rest of the gradient due to its milder conditions. Earlywood and latewood were unrelated at the three upper sites, and both compartments responded to different climatic factors. Hydraulic conductivity determined by vessel size was used to describe tree performance, which was modulated by temperature along the gradient. We hypothesize that the main processes involved are the timing of earlywood formation and carbohydrate dynamics. Water availability during late spring and summer affected latewood width, but only at low elevation.This paper illustrates the complexity of analyzing climate-growth relationships in oceanic areas under the absence of a prevailing limiting factor, while providing a feasible explanation of potential mechanisms involved.     

Seasonal conductivity and embolism in the roots and stems of two clonal ring-porous trees, Sassafras albidum (Lauraceae) and Rhus typhina (Anacardiaceae)

Seasonal xylem (wood) conductivity and embolism (air blockage) patterns were monitored in roots vs. stems of two clonal ring-porous tree species, Sassafras albidum and Rhus typhina, throughout 1996 and 1997. Stems of both species were 100% embolized in the early spring and became conductive by late June following leaf expansion and maturation of new earlywood vessels. Dyes indicated the stem conduction was restricted almost exclusively to the current year's growth ring. Stems became totally embolized again by early October, before the first freezing temperatures. In contrast, woody roots of both species maintained low embolism values, many conductive growth rings, and high conductivity values regardless of the season. No positive root pressures were detected in either species. The mean frost depth (204 ± 11 mm) was deeper than all sampled roots of Rhus and 47% of sampled roots of Sassafras. The roots that had been in frozen soil either avoided embolism altogether or they were able to reverse embolism by a mechanism other than positive root pressures.     

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

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