Earlywood Vessels Research Articles

Seasonal activity and phoresy rates of Nitidulid beetles (Coleoptera: Nitidulidae) captured in stands with oak wilt infections in northern Michigan, USA.

Nitidulid beetles (Coleoptera: Nitidulidae), the overland vectors of the Bretziella fagacearum fungus that causes oak wilt, were monitored in infection centers in Quercus rubra stands in northern Michigan, USA using baited, wind-oriented traps for 2 years. First nitidulid captures, accounting for <1.5% of total annual captures, occurred in late April in both years (<50 cumulative degree days [DDs]; base 10°C). A total of 1,188 and 735 beetles representing 19 and 18 species were captured in 2018 (18 traps; 3 sites) and 2019 (16 traps; 4 sites), respectively. Ground traps (1.5 m high) captured more beetles and species than traps on canopy branches of red oaks. Most nitidulids (81-86%) were captured in May and June, but frequent precipitation and cool spring temperatures extended activity into early July in 2019. In 2018, 336 beetles representing 12 species were screened for B. fagacearum spores, but only 20 beetles from 4 species bore viable spores. Mycelial mats on red oaks killed were 4-fold more abundant in 2019 than in 2018. Of the 225 beetles screened in 2019, 56 beetles representing 6 species had viable spores. Nearly all (96%) spore-bearing beetles in both years were captured in late May or June. In bimonthly xylem samples collected from healthy trees, large earlywood vessels, presumably more vulnerable to infection than latewood, were present from May to June in 2018 and until early July in 2019. Results are consistent with mid-May to mid-July high-risk periods designated in current state and regional guidelines for oak wilt management.

Responses to drought of two Mediterranean ring-porous, deciduous species: Searching for climate smart trees and shrubs

Drought-tolerant tree species with high growth rates and a good capacity for carbon storage in woody tissues (dense wood) are searched for due to aridification. Deciduous, ring-porous tree and shrub species could show such drought tolerance and growth traits, thus representing good candidates for climate-smart rewilding. However, we still do not know the long-term growth rates of these species and how they respond to drought, particularly in climate change hotspots such as the Mediterranean Basin. We analysed these issues at the site and individual levels in two ring-porous, deciduous species (Pistacia terebinthus, Celtis australis) using dendroecology and wood anatomy. The ring width, earlywood vessel diameter, vessel density (VD) and area (%) were measured in two focal sites, one per species, and then growth data were compared with two secondary sites to test if site-to-site synchrony changed through time. Ring-width indices (RWI) and the hydraulic diameter (Dh) of earlywood were calculated. Growth rates (ring width), Dh and vessel area were higher in C. australis (1.03−2.26 mm, 269 μm, 33.9 %) than in P. terebinthus (0.57−0.72 mm, 146 μm, 21.5 %). Consequently, VD was higher in P. terebinthus than in C. australis (104 vs. 61 vessels mm−2). The ring width and Dh were more coupled in P. terebinthus (r = 0.43) than in C. australis (r = 0.32). RWI series of the focal and secondary sites have been synchronized since the 1990s as temperatures rose. Precipitation during the growing season (May, June) enhanced growth and VD of both species. P. terebinthus was more responsive to a drought index than C. australis. The two study species show high growth rates and tolerate drought being thus suitable candidates for climate-smart rewilding.

Performance of convolutional neural network (CNN) and performance influencing factors for wood species classification of Lepidobalanus growing in Korea

This study aimed to investigate the performance and factors affecting the species classification of convolutional neural network (CNN) architecture using whole-part and earlywood-part cross-sectional datasets of six Korean Quercus species. The accuracy of species classification for each condition was analyzed using the datasets, data augmentation, and optimizers—stochastic gradient descent (SGD), adaptive moment estimation (Adam), and root mean square propagation (RMSProp)—based on a CNN architecture with three to four convolutional layers. The model trained with the augmented dataset yielded significantly superior results in terms of classification accuracy compared to the model trained with the non-augmented dataset. The augmented dataset was the only factor affecting classification accuracy in the final five epochs. In contrast, four factors in the entire epoch, such as the Adam and SGD optimizers and the earlywood-part and whole-part datasets, affected species classification. The arrangement of earlywood vessels, broad rays, and axial parenchyma was identified as a major influential factor in the CNN species classification using gradient-weighted class activation mapping (Grad-CAM) analysis. The augmented whole-part dataset with the Adam optimizer achieved the highest classification accuracy of 85.7% during the final five epochs of the test phase.

Origin-specific differences in the durability of black locust (Robinia pseudoacacia) wood against wood-destroying basidiomycetes

Global climate change is accompanied by a change in tree composition in many regions. In Europe, the distribution areas of many species are expanding towards the north so that, among others, black locust (Robinia pseudoacacia), which is native to the USA and has long been established in south-eastern Europe, is also becoming increasingly important in central and northern Europe. Many other tree species are known to have different properties between their original and new locations, including the biological durability of the wood. Hence, the resistance of black locust wood against decay fungi was studied concerning origin-specific differences. Wood was sampled from seven different origins in Europe and original habitats in the United States. Fungal incubation experiments were conducted, wood extractives were analysed, and different anatomical characteristics were quantified such as ring width, vessel size distribution and the presence of tyloses. In addition to differences in durability between juvenile and mature wood, origin-specific differences within the mature heartwood were attributed to extractive contents and the percentages of earlywood vessels containing tyloses. Based on parameters that contributed at least 20% to mass loss, susceptibility to fungal decay was modelled with multiple regressions.

Ring- and diffuse-porous tree species coexisting in cold and humid temperate forest diverge in stem and leaf phenology

Leaf and stem phenology are critical drivers of tree growth patterns in seasonal climates, but the implications for species differences in radial stem growth dynamics are still poorly studied. In our study, we compared the leaf phenology and stem phenology with the underlying cell development as well as annual stem growth between five diffuse-porous (DP) and five ring-porous (RP) coexisting angiosperm species in cold and humid temperate forests. Our results show that RP species unfolded leaves later but initiated wood growth earlier compared to DP species. Yet, xylem vessel maturation in the stem started in June and was remarkably synchronized between DP and RP, which implies that species from both functional groups can effectively avoid vessel cavitation potentially triggered by late spring frost. DP species exhibited one peak in stem growth across the growing season reflecting a uniform vessel formation pattern. Instead, RP species exhibited two peaks in stem growth, with the first peak reflecting expansion of early-wood vessels and the second peak related to subsequent fiber and vessel proliferation in the late-wood. In general, species with a greater number of growth days from the start of cambium activity until full lignification of cells exhibited higher annual stem growth, regardless of species group. The observed differences in leaf and stem phenology between DP and RP species are discussed with respect to the adaptation potential of the two functional groups to changing climate conditions in cold and humid temperate forests.

Radial Variation of Wood Anatomical Characteristics and Maturation Ages of Six Korean Oak Species

The objective of this study was to examine and compare radial variation of the anatomical characteristics and the transition age from juvenile to mature wood of the six Korean oak wood species: Quercus variabilis (Qv), Quercus serrata (Qs), Quercus mongolica (Qm), Quercus dentata (Qd), Quercus aliena (Qal), and Quercus acutissima (Qac). Quantitative anatomical features were observed from the pith to the bark at five growth-ring intervals using optical microscopy. A segmented regression model was used to evaluate the transition from juvenile wood to mature wood. The clearest transition from juvenile to mature wood was observed in the radial variation of the earlywood vessel diameter and fiber length. The maturation age of the six Korean oak species ranged from 19 to 44 years. Qv exhibited the highest values for latewood vessel diameter, fiber length, and fiber diameter. Qac displayed the highest values for earlywood vessel diameter and fiber wall thickness. The highest fiber lumen diameter was observed for Qm. The differences in earlywood and latewood vessel diameters and fiber lengths could be utilized as identification keys for these species. Earlywood vessel diameter and fiber length have emerged as the most reliable indicators for estimating the transition from juvenile to mature wood.

No effect of snow on shrub xylem traits: Insights from a snow-manipulation experiment on Disko Island, Greenland

Widespread shrubification across the Arctic has been generally attributed to increasing air temperatures, but responses vary across species and sites. Wood structures related to the plant hydraulic architecture may respond to local environmental conditions and potentially impact shrub growth, but these relationships remain understudied. Using methods of dendroanatomy, we analysed shrub ring width (RW) and xylem anatomical traits of 80 individuals of Salix glauca L. and Betula nana L. at a snow manipulation experiment in Western Greenland. We assessed how their responses differed between treatments (increased versus ambient snow depth) and soil moisture regimes (wet and dry). Despite an increase in snow depth due to snow fences (28–39 %), neither RW nor anatomical traits in either species showed significant responses to this increase. In contrast, irrespective of the snow treatment, the xylem specific hydraulic conductivity (Ks) and earlywood vessel size (LA95) for the study period were larger in S. glauca (p < 0.1, p < 0.01) and B. nana (p < 0.01, p < 0.001) at the wet than the dry site, while both species had larger vessel groups at the dry than the wet site (p < 0.01). RW of B. nana was higher at the wet site (p < 0.01), but no differences were observed for S. glauca. Additionally, B. nana Ks and LA95 showed different trends over the study period, with decreases observed at the dry site (p < 0.001), while for other responses no difference was observed. Our results indicate that, taking into account ontogenetic and allometric trends, hydraulic related xylem traits of both species, along with B. nana growth, were influenced by soil moisture. These findings suggest that soil moisture regime, but not snow cover, may determine xylem responses to future climate change and thus add to the heterogeneity of Arctic shrub dynamics, though more long-term species- and site- specific studies are needed.

Flood rings, earlywood vessels and hydrological signal in Fraxinus pennsylvanica trees growing along the central Assiniboine river floodplain, southcentral Canada

In southcentral Canada, paleoflood reconstructions have mainly focussed on identifying flood rings in upper terrace bur oak (Quercus macrocarpa) trees from the Red and Assiniboine rivers. In contrast to upper terrace trees, floodplain tree species such as green ash (Fraxinus pennsylvanica) were said a few decades ago to provide ‘noisy’ flood proxies due to a greater flood exposition. This unverified assumption about floodplain trees was tested using green ash trees growing along the floodplain of the Assiniboine river. In each sample and for each year, flood rings were visually identified, vessels with a cross-sectional area >1000 µm2 as well as earlywood, latewood and total ring width were measured. Results indicated that flood rings were replicable and that they correspond to tree rings characterized by a high number of earlywood vessels having a reduced mean area (less porous earlywood) and, in which, radial multiples were abundant. Years in which flood rings were most abundant corresponded to documented high-magnitude floods. Flood-ring, earlywood vessel (except density and total area) and ring-width chronologies were all significantly correlated to spring mean discharge. Floodplain green ash trees do capture hydrological events including extreme floods and should thus not be ignored in paleoflood studies.

Time-moisture superposition principle in creep behavior of white oak with various earlywood vessel locations

Creep behavior of wood plays a fundamental role in precision processing of wood. In this work, experimental creep tests have been conducted to determine the influence of earlywood vessel location and moisture content on creep behavior of Quercus alba (white oak). Time-moisture superposition principle was applied to predict long-term creep behavior of white oak. Results revealed that both of instantaneous and 45-min strain of specimens increased with the increasing of moisture content and decreased with increasing distance between earlywood vessel belt and load-bearing surface significantly. Additionally, the time-moisture superposition principle was found to have feasibility to predict creep behavior of white oak with various earlywood vessel locations and moisture content ranges (6 % - 18 %). We believe that the proposed investigation was beneficial for the processing precision and civil engineering applications of wood.

Anatomical Characteristics and Fibre Quality of Grapevine (Vitis vinifera L.) Stem Wood

This study investigated the anatomical characteristics and fibre quality for papermaking indices of Vitis vinifera L. (grapevine) stem wood, which is extracted as agricultural waste. Two grapevine trunks were collected from the G&amp;uuml;lnar region in Turkey. Observations on microscopic anatomical characteristics were carried out on sectioned and macerated wood samples. According to the measurements conducted, the following mean anatomical characteristics were determined: earlywood vessel tangential diameter 258.81 &amp;mu;m, latewood vessel tangential diameter 35.52 &amp;mu;m, ray width 197.19 &amp;mu;m, ray height 4618.67 &amp;mu;m, vessel length 498.85 &amp;mu;m, fibre length 1.03 mm, fibre diameter 22.05 &amp;mu;m, and fibre wall thickness 4.23 &amp;mu;m. Based on the determined characteristics linked to the fibre quality, the fibres of the grapevine can be placed in Quality Class III for pulp and paper processing. All derived indices of grapevine met the acceptable threshold except for the flexibility ratio. Examining the anatomical structure of the grapevine will enable a database to be created for further studying of wood anatomy and these characteristics can be evaluated with respect to other possible areas of use.

Structural Variation Patterns in Xylem Vessels and Parenchyma Cells and Their Association with Tree Evolution

Xylem vessels and parenchyma cells perform functions such as water transport and nutrient storage in trees. However, they are highly variable in different trees. Therefore, this study aimed to explore the structural change patterns in vessels and parenchyma cells in the sapwood, transition wood, and heartwood of tree species with different degrees of evolution. The structural characteristics of the two types of cells in the sapwood, transitional wood, and heartwood were measured in six species with different levels of evolution, namely, Michelia macclurei Dandy, Cinnamomum camphora (L.) presl, Erythrophleum fordii Oliv, Melaleuca leucadendron L., Parashorea chinensis Wang Hsie and Tectona grandis L.F. The results showed that the more evolved species had larger earlywood vessel lumen diameters, thicker walls, and wider hydraulic diameters, as well as smaller latewood vessel densities, reflecting better water transport effectiveness and higher safety. From the sapwood to the transition wood and heartwood, the earlywood vessel lumen diameter of the more primitive species tended to be stable and then decrease, while that of the more evolved species gradually decreased. The latewood vessel density of the more primitive species tended to be stable and then increase, while that of the more evolved species tended to be stable. Additionally, the starch grains of the more primitive species were mainly distributed in the axial or ray parenchyma cells of the sapwood, while those of the more evolved species were abundantly distributed in the axial and ray parenchyma cells of the sapwood, and the distribution of the starch grains was reduced in the transition wood and heartwood. From the sapwood to the heartwood, the ray parenchyma cell fraction tended to be stable in the more primitive species, and tended to be stable before decreasing in the more evolved species. The pit density in the horizontal wall of the ray parenchyma cells tended to be stable and decrease in the more primitive species, while in the more evolved species it tended to be stable or decrease before stabilizing. Overall, trees’ vessels have gradually undergone the optimal selection of vessels during evolution, and the structural variation in the parenchyma cells contributes to their nutrient storage and transport.

Earlywood Anatomy Highlights the Prevalent Role of Winter Conditions on Radial Growth of Oak at Its Distribution Boundary in NW Iberia

We compared climate-growth relationships (1956-2013) of two natural pedunculate oak (Quercus robur L.) stands with different water-holding capacities growing at the species distribution limit of the Mediterranean Region in NW Iberia. For this, tree-ring chronologies of earlywood vessel size (separating the first row from the other vessels) and latewood width were obtained. Earlywood traits were coupled to conditions during dormancy, whereby an elevated winter temperature appears to induce a high consumption of carbohydrates, resulting in smaller vessels. This effect was reinforced by waterlogging at the wettest site, whose correlation to winter precipitation was strongly negative. Soil water regimes caused differences between vessel rows, since all earlywood vessels were controlled by winter conditions at the wettest site, but only the first row at the driest one; radial increment was related to water availability during the previous rather than the current season. This confirms our initial hypothesis that oak trees near their southern distribution boundary adopt a conservative strategy, prioritizing reserve storage under limiting conditions during the growing period. We believe that wood formation is highly dependent on the balance between the previous accumulation of carbohydrates and their consumption to maintain both respiration during dormancy and early spring growth.

Revue documentaire et nouvelles perspectives de l’anatomie du bois aidant à mieux comprendre et lutter contre le flétrissement du chêne

Oak wilt, which develops in the outer sapwood, is the most destructive disease of oaks in the United States. Species of red oaks are more susceptible to this disease than white oak species and are more likely to facilitate the spread of the pathogen Bretziella fagacearum. To prevent its establishment in new areas, phytosanitary certificates are mandatory for commercial trade, as is the inspection of logs to confirm identification. A literature survey and the results of our assays with seven oak species confirm that it is easy to identify wood of the red and white oak groups using anatomical features. Specifically, earlywood vessels are generally open in red oaks, while they appear occluded with tyloses in white oaks. Such plugs are the consequence of air embolism (cavitation) and not the cause of the wilting process. Although these wide vessels are efficient for water transport, they are vulnerable to cavitation that appears to favour the growth of the oak wilt pathogen. Compartmentalization of infected wood succeeds in restraining cavitation and pathogen spread, allowing some trees to recover from oak wilt.

Linking xylem structure and function: the comparative method in from the cold.

This article is a Commentary on Savage et al. (2022), 235: 953–964.

Different xylogenesis responses to atmospheric water demand contribute to species coexistence in a mixed pine–oak forest

Seasonal patterns of wood formation (xylogenesis) remain understudied in mixed pine–oak forests despite their contribution to tree coexistence through temporal niche complementarity. Xylogenesis was assessed in three pine species (Pinus cembroides, Pinus leiophylla, Pinus engelmannii) and one oak (Quercus grisea) coexisting in a semi-arid Mexican forest. The main xylogenesis phases (production of cambium cells, radial enlargement, cell-wall thickening and maturation) were related to climate data considering 5–15-day temporal windows. In pines, cambium activity maximized from mid-March to April as temperature and evaporation increased, whereas cell radial enlargement peaked from April to May and was constrained by high evaporation and low precipitation. Cell-wall thickening peaked from June to July and in August–September as maximum temperature and vapour pressure deficit (VPD) increased. Maturation of earlywood and latewood tracheids occurred in May–June and June–July, enhanced by high minimum temperatures and VPD in P. engelmannii and P. leiophylla. In oak, cambial onset started in March, constrained by high minimum temperatures, and vessel radial enlargement and radial increment maximized in April as temperatures and evaporation increased, whereas earlywood vessels matured from May to June as VPD increased. Overall, 15-day wet conditions enhanced cell radial enlargement in P. leiophylla and P. engelmannii, whereas early-summer high 15-day temperature and VPD drove cell-wall thickening in P. cembroides. Warm night conditions and high evaporation rates during spring and summer enhanced growth. An earlier growth peak in oak and a higher responsiveness to spring–summer water demand in pines contributed to their coexistence.

Using tree-ring width and earlywood vessel features to study the decline of Quercus brantii Lindl in Zagros forests of Iran

Using tree-ring width and earlywood vessel features to study the decline of Quercus brantii Lindl in Zagros forests of Iran

Seasonal patterns of increases in stem girth, vessel development, and hydraulic function in deciduous tree species.

The onset of spring growth and vessel formation were examined within three deciduous woody plant species, Acer rubrum, Populus balsamifera ssp. trichocarpa and Quercus rubra. We were broadly interested in the lag between the onset of girth expansion and the formation of mature and hydraulically conductive vessels within the new xylem. Dendrometers were installed on 20 trees (6-7 per species), and expansion of both bole and distal stems was monitored throughout the growing season in a common garden. For each species, four to six distal stems were harvested every other week for anatomical examination of vessel formation. Additionally, for Populus and Quercus, hydraulic conductivity measurements and active xylem staining were completed on all stem samples. For all three species, the timing of girth expansion was similar. Expansion of distal branches occurred 12-37 d earlier than that of the bole. Vessel formation initiated several weeks prior to leaf-out, but no new earlywood vessels were mature at the time of bud break for Acer and Populus and only a few were present in Quercus. Initial stem girth expansion occurred 2 to &gt;6 weeks before the maturation of the first current-year vessels, and there was an additional delay of up to 4 weeks before mature vessels became hydraulically functional. Hydraulic conductivity was strongly correlated with the number and diameter of stained vessels. Bud break and leaf expansion relied predominantly on water supplied by vessels formed during prior seasons. Early-season activity is likely affected by the function of older xylem vessels and the environmental factors that influence their structure and function. Understanding the functional lifespan of vessels and the varying contributions of new and older vessels to conductivity are critical to understanding of the phenology and vascular function of long-lived woody plants in response to changing climates.

Different Responses in Vascular Traits between Dutch Elm Hybrids with a Contrasting Tolerance to Dutch Elm Disease.

The ascomycetous fungus Ophiostoma novo-ulmi is the causative agent of the current Dutch elm disease (DED) pandemic, which has ravaged many tens of millions of European and North American elm trees. Host responses in vascular traits were studied in two Dutch elm hybrids, ‘Groeneveld’ and ‘Dodoens’, which show different vascular architecture in the secondary xylem and possess contrasting tolerances to DED. ‘Groeneveld’ trees, sensitive to DED, possessed a high number of small earlywood vessels. However, these trees showed a poor response to DED infection for the earlywood vascular characteristics. Following infection, the proportion of least vessels with a vessel lumen area less than 2500 µm2 decreased from 65.4% down to 53.2%. A delayed response in the increasing density of vessels showing a reduced size in the latewood prevented neither the rapid fungal spread nor the massive colonisation of the secondary xylem tissues resulting in the death of the infected trees. ‘Dodoens’ trees, tolerant to DED, possessed a low number of large earlywood vessels and showed a prominent and fast response to DED infection. Vessel lumen areas of newly formed earlywood vessels were severely reduced together with the vessel size : number ratio. Following infection, the proportion of least vessels with a vessel lumen area less than 2500 µm2 increased from 75.6% up to 92.9%. A trend in the increasing density of vessels showing a reduced size was maintained not only in the latewood that was formed in the year of infection but also in the earlywood that was formed in the consecutive year. The occurrence of fungal hyphae in the earlywood vessels that were formed a year following the infection was severely restricted, as revealed by X-ray micro-computed tomography imaging. Possible reasons responsible for a contrasting survival of ‘Groeneveld’ and ‘Dodoens’ trees are discussed.

Leaf photosynthetic capacity, trunk wood structure and stem xylem sap flow in 700-years old Quercus robur L.: a pilot study upon oak ‘Bartek’, a natural monument in Poland

Physiological studies of long-lived trees are particularly important at this time, especially in light of the need for trees to adapt to global climate change. The results of the present studies were obtained on an approximately 700-year-old Quercus robur L. – the ‘Bartek’ oak. The tree has to adapt to changing climatic conditions, starting from the transition between the Medieval Warm Period and the Little Ice Age, up to the present time of rapid global climate change. Tomograph imaging showed decay of the tree trunk interior and revealed that undamaged wood forms a thin layer around the trunk perimeter. Two series of experiments were carried out to assess the physiological state of the tree. The first concerned measurements related to photosynthetic capacity: chlorophyll a fluorescence, gas exchange (CO2 assimilation, transpiration), stomatal conductance and leaf water potential. The second series concerned xylem sap flow velocity and anatomical studies of stem wood. Photosynthetic capacity was within the limits reported for young healthy trees. The diurnal pattern of velocity of xylem sap flow was also typical for young vigorous trees and flow velocity correlated positively with solar radiation and negatively with air relative humidity. Anatomical observations of the outermost wood showed relatively narrow annuals rings with large diameter earlywood vessels. The results indicate that the veteran tree does not show signs of water stress probably due to a good balance of water flow and that leaf area of the canopy needs only the current ring of wood to feed transpiration of the canopy.

Effects of nitrogen addition and increased precipitation on xylem growth of Quercus acutissima Caruth. in central China.

Atmospheric nitrogen (N) deposition and increasing precipitation affect carbon sequestration in terrestrial ecosystems, but how these two concurrent global change variables affect xylem growth in trees (i.e., independently or interactively) remains unclear. We conducted novel experiments in central China to monitor the xylem growth in a dominant species (Quercus acutissima Caruth.) in response to N addition (CN), supplemental precipitation (CW) or both treatments (CNW), compared with untreated controls (C). Measurements were made at weekly intervals during 2014-15. We found that supplemental precipitation significantly enhanced xylem growth in the dry spring of 2015, indicating a time-varying effect of increased precipitation on intra-annual xylem growth. Elevated N had no significant effect on xylem increment, xylem growth rate, and lumen diameters and potential hydraulic conductivity (Ks) of earlywood vessels, but Ks with elevated N was significantly negatively related to xylem increment. The combination of additional N and supplemental precipitation suppressed the positive effect of supplemental precipitation on xylem increment in the dry spring of 2015. These findings indicated that xylem width was more responsive to supplemental precipitation than to increasing N in a dry early growing season; the positive effect of supplemental precipitation on xylem growth could be offset by elevated N resources. The negative interactive effect of N addition and supplemental precipitation also suggested that increasing N deposition and precipitation in the future might potentially affect carbon sequestration of Q. acutissima during the early growing season in central China. The effects of N addition and supplemental precipitation on tree growth are complex and might vary depending on the growth period and local climatic conditions. Therefore, future models of tree growth need to consider multiple-time scales and local climatic conditions when simulating and projecting global change.