Wood specific gravity and anatomy in Heliocarpus appendiculatus (Tiliaceae)

Specific gravity exhibits extremely large radial increases with distance from the pith in Heliocarpus appendiculatus Turcz. (Tiliaceae), a pioneer of neotropical wet forests. To determine some of the wood anatomical changes associated with this increase, wood samples taken at breast height from three trees were divided into 1.0-cm-long segments from pith to bark. Measurements were made of fiber wall thickness, fiber lumen diameter, and percentages of fibers, axial parenchyma, ray parenchyma, and vessels on sections prepared from each segment. The extreme radial increases in specific gravity were associated with increases in fiber wall thickness, decreases in fiber diameter, decreases in fiber lumen diameter, and changes in the relative proportions of fibers and parenchyma. The increase in percent fiber concomitant with a decrease in axial parenchyma was the most important contributor to the increase in specific gravity in this species. The best predictor of specific gravity was percent fibers (r = 0.91, 0.92, 0.94) or percent axial parenchyma (r = -0.92, -0.91, -0.95), two variables that were highly intercorrelated (r = -0.95, -0.98, -0.99).     

Wood anatomy and carbon‐isotope discrimination support long‐term hydraulic deterioration as a major cause of drought‐induced dieback

Hydraulic impairment due to xylem embolism and carbon starvation are the two proposed mechanisms explaining drought‐induced forest dieback and tree death. Here, we evaluate the relative role played by these two mechanisms in the long‐term by quantifying wood‐anatomical traits (tracheid size and area of parenchyma rays) and estimating the intrinsic water‐use efficiency (iWUE) from carbon isotopic discrimination. We selected silver fir and Scots pine stands in NE Spain with ongoing dieback processes and compared trees showing contrasting vigour (declining vs nondeclining trees). In both species earlywood tracheids in declining trees showed smaller lumen area with thicker cell wall, inducing a lower theoretical hydraulic conductivity. Parenchyma ray area was similar between the two vigour classes. Wet spring and summer conditions promoted the formation of larger lumen areas, particularly in the case of nondeclining trees. Declining silver firs presented a lower iWUE than conspecific nondeclining trees, but the reverse pattern was observed in Scots pine. The described patterns in wood anatomical traits and iWUE are coherent with a long‐lasting deterioration of the hydraulic system in declining trees prior to their dieback. Retrospective quantifications of lumen area permit to forecast dieback in declining trees 2–5 decades before growth decline started. Wood anatomical traits provide a robust tool to reconstruct the long‐term capacity of trees to withstand drought‐induced dieback.     

木本植物木质部栓塞脆弱性研究新进展

木质部空穴化和栓塞是木本植物在干旱等条件下遭受水分胁迫时产生的木质部输水功能障碍, 在全球气候变化的大背景下, 栓塞脆弱性对干旱响应的研究已成为热点和重要内容。近年来有关木质部栓塞脆弱性与植物输水结构和耐旱性的关系已有大量研究并取得一定成果, 但是, 不同学者在不同地区对不同材料的研究结果存在很大不同。该文就近年来这一研究领域取得的成果及争议问题进行了概括和总结, 主要涉及木质部栓塞脆弱性(P₅₀)及脆弱曲线的建立方法、木质部栓塞脆弱性与木质部结构(导管直径、导管长度、纹孔膜、木质部密度、纤维及纤维管胞)间的关系和木质部栓塞脆弱性与耐旱性的关系, 并对未来工作进行展望, 提出在未来的工作中应对同一树种使用Cochard Cavitron离心机法、Sperry离心机技术与传统方法建立的脆弱曲线进行比较验证、计算P₅₀值、分析植物个体器官水平差异(根、茎、叶)、测定树种生理生态指标, 探索植物栓塞脆弱性与输水结构和耐旱性的关系, 从而评估不同类型植物在未来气候变化下的耐旱能力。     

A review of new research progress on the vulnerability of xylem embolism of woody plants

Xylem cavitation/embolism is the blockage of xylem conduits when woody plants suffer from water stress under drought and other environmental conditions, the study of embolism has become a hot and key topic under global climate change. Recent researches on the relationship between the vulnerability of xylem embolism and hydraulic architecture/drought tolerance have made some progress, however, scholars reached different conclusions based on results from different regions or different materials. This paper reviews the current achievements and controversial viewpoints, which includes indicator of xylem embolism vulnerability (P₅₀), method of vulnerability curve establishment, the relationship between embolism vulnerability and hydraulic architecture (vessel diameter, vessel length, pit area, wood density, fiber and fiber tracheid) and the relationship between embolism vulnerability and drought tolerance of woody plants. Future studies should use Cochard Cavitron centrifuge and Sperry centrifuge coupled with traditional methods to establish vulnerability curves, calculate P₅₀, analyze the difference among different organisms (root, stem, leaf), and measure physiological and ecological indexes. Future studies should be aimed to explore the relationship between the vulnerability of xylem embolism and hydraulic architecture/drought tolerance and to assess drought tolerance ability of different species under future climate change.     

Amazonia trees have limited capacity to acclimate plant hydraulic properties in response to long‐term drought

The fate of tropical forests under future climate change is dependent on the capacity of their trees to adjust to drier conditions. The capacity of trees to withstand drought is likely to be determined by traits associated with their hydraulic systems. However, data on whether tropical trees can adjust hydraulic traits when experiencing drought remain rare. We measured plant hydraulic traits (e.g. hydraulic conductivity and embolism resistance) and plant hydraulic system status (e.g. leaf water potential, native embolism and safety margin) on >150 trees from 12 genera (36 species) and spanning a stem size range from 14 to 68 cm diameter at breast height at the world's only long‐running tropical forest drought experiment. Hydraulic traits showed no adjustment following 15 years of experimentally imposed moisture deficit. This failure to adjust resulted in these drought‐stressed trees experiencing significantly lower leaf water potentials, and higher, but variable, levels of native embolism in the branches. This result suggests that hydraulic damage caused by elevated levels of embolism is likely to be one of the key drivers of drought‐induced mortality following long‐term soil moisture deficit. We demonstrate that some hydraulic traits changed with tree size, however, the direction and magnitude of the change was controlled by taxonomic identity. Our results suggest that Amazonian trees, both small and large, have limited capacity to acclimate their hydraulic systems to future droughts, potentially making them more at risk of drought‐induced mortality.     

Are lianas more drought-tolerant than trees? A test for the role of hydraulic architecture and other stem and leaf traits

Lianas are an important component of Neotropical forests, where evidence suggests that they are increasing in abundance and biomass. Lianas are especially abundant in seasonally dry tropical forests, and as such it has been hypothesized that they are better adapted to drought, or that they are at an advantage under the higher light conditions in these forests. However, the physiological and morphological characteristics that allow lianas to capitalize more on seasonal forest conditions compared to trees are poorly understood. Here, we evaluate how saplings of 21 tree and liana species from a seasonal tropical forest in Panama differ in cavitation resistance (P ₅₀) and maximum hydraulic conductivity (K _h), and how saplings of 24 tree and liana species differ in four photosynthetic leaf traits (e.g., maximum assimilation and stomatal conductance) and six morphological leaf and stem traits (e.g., wood density, maximum vessel length, and specific leaf area). At the sapling stage, lianas had a lower cavitation resistance than trees, implying lower drought tolerance, and they tended to have a higher potential hydraulic conductivity. In contrast to studies focusing on adult trees and lianas, we found no clear differences in morphological and photosynthetic traits between the life forms. Possibly, lianas and trees are functionally different at later ontogenetic stages, with lianas having deeper root systems than trees, or experience their main growth advantage during wet periods, when they are less vulnerable to cavitation and can achieve high conductivity. This study shows, however, that the hydraulic characteristics and functional traits that we examined do not explain differences in liana and tree distributions in seasonal forests.     

Hydraulic failure and tree dieback are associated with high wood density in a temperate forest under extreme drought

Catastrophic hydraulic failure will likely be an important mechanism contributing to large‐scale tree dieback caused by increased frequency and intensity of droughts under global climate change. To compare the susceptibility of 22 temperate deciduous tree and shrub species to hydraulic failure during a record drought in the southeastern USA, we quantified leaf desiccation, native embolism, wood density, stomatal conductance and predawn and midday leaf water potential at four sites with varying drought intensities. At the two driest sites, there was widespread leaf wilting and desiccation, and most species exhibited predawn leaf water potentials of ≤3 MPa and >60% loss of xylem conductivity in branches. Although species with high wood density were more resistant to cavitation, they had higher levels of native embolism and greater canopy dieback than species with low wood density. This unexpected result can be explained by the failure of species with dense wood to avert a decline in water potential to dangerous levels during the drought. Leaf water potential was negatively correlated with wood density, and the relationship was strongest under conditions of severe water deficit. Species with low wood density avoided catastrophic embolism by relying on an avoidance strategy that involves partial drought deciduousness, higher sensitivity of stomata to leaf water potential and perhaps greater rooting depth. These species therefore maintained water potential at levels that ensured a greater margin of safety against embolism. These differences among species may mediate rapid shifts in species composition of temperate forests if droughts intensify due to climate change.     

Vulnerability to xylem embolism as a major correlate of the environmental distribution of rain forest species on a tropical island

Increases in drought‐induced tree mortality are being observed in tropical rain forests worldwide and are also likely to affect the geographical distribution of tropical vegetation. However, the mechanisms underlying the drought vulnerability and environmental distribution of tropical species have been little studied. We measured vulnerability to xylem embolism (P₅₀) of 13 woody species endemic to New Caledonia and with different xylem conduit morphologies. We examined the relation between P₅₀, along with other leaf and xylem functional traits, and a range of habitat variables. Selected species had P₅₀ values ranging between ?4.03 and ?2.00 MPa with most species falling in a narrow range of resistance to embolism above ?2.7 MPa. Embolism vulnerability was significantly correlated with elevation, mean annual temperature and percentage of species occurrences located in rain forest habitats. Xylem conduit type did not explain variation in P₅₀. Commonly used functional traits such as wood density and leaf traits were not related to embolism vulnerability. Xylem embolism vulnerability stands out among other commonly used functional traits as a major driver of species environmental distribution. Drought‐induced xylem embolism vulnerability behaves as a physiological trait closely associated with the habitat occupation of rain forest woody species.     

Xylem hydraulic safety and construction costs determine tropical tree growth

Faster growth in tropical trees is usually associated with higher mortality rates, but the mechanisms underlying this relationship are poorly understood. In this study, we investigate how tree growth patterns are linked with environmental conditions and hydraulic traits, by monitoring the cambial growth of 9 tropical cloud forest tree species coupled with numerical simulations using an optimization model. We find that fast‐growing trees have lower xylem safety margins than slow‐growing trees and this pattern is not necessarily linked to differences in stomatal behaviour or environmental conditions when growth occurs. Instead, fast‐growing trees have xylem vessels that are more vulnerable to cavitation and lower density wood. We propose the growth ‐ xylem vulnerability trade‐off represents a wood hydraulic economics spectrum similar to the classic leaf economic spectrum, and show through numerical simulations that this trade‐off can emerge from the coordination between growth rates, wood density, and xylem vulnerability to cavitation. Our results suggest that vulnerability to hydraulic failure might be related with the growth‐mortality trade‐off in tropical trees, determining important life history differences. These findings are important in furthering our understanding of xylem hydraulic functioning and its implications on plant carbon economy.     

Intraspecific variation in embolism resistance and stem anatomy across four sunflower (Helianthus annuus L.) accessions

Drought‐induced xylem embolism is a key process closely related to plant mortality during extreme drought events. However, this process has been poorly investigated in crop species to date, despite the observed decline of crop productivity under extreme drought conditions. Interspecific variation in hydraulic traits has frequently been reported, but less is known about intraspecific variation in crops. We assessed the intraspecific variability of embolism resistance in four sunflower (Helianthus annuus L.) accessions grown in well‐watered conditions. Vulnerability to embolism was determined by the in situ flow‐centrifuge method (cavitron), and possible trade‐offs between xylem safety, xylem efficiency and growth were assessed. The relationship between stem anatomy and hydraulic traits was also investigated. Mean P ₅₀ was ?3 MPa, but significant variation was observed between accessions, with values ranging between ?2.67 and ?3.22 MPa. Embolism resistance was negatively related to growth and positively related to xylem‐specific hydraulic conductivity. There is, therefore, a trade‐off between hydraulic safety and growth but not between hydraulic safety and efficiency. Finally, we found that a few anatomical traits, such as vessel density and the area of the vessel lumen relative to that of the secondary xylem, were related to embolism resistance, whereas stem tissue lignification was not. Further investigations are now required to investigate the link between the observed variability of embolism resistance and yield, to facilitate the identification of breeding strategies to improve yields in an increasingly arid world.     

Seasonal development of secondary xylem and phloem in <Emphasis Type="Italic">Schizolobium parahyba</Emphasis> (Vell.) Blake (Leguminosae: Caesalpinioideae)

The cambial activity and periodicity of secondary xylem and phloem formation have been less studied in tropical tree species than in temperate ones. This paper describes the relationship between seasonal cambial activity, xylem and phloem development, and phenology in Schizolobium parahyba, a fast growing semideciduous seasonal forest tree from southeastern Brazil. From 2002 to 2003, wood samples were collected periodically and phenology and climate were recorded monthly in the same period. S. parahyba forms annual growth increments in wood, delimited by narrow initial parenchyma bands. The reduction of the cambial activity to a minimum correlates to the dry season and leaf fall. The higher cambial activity correlates to the wet season and the presence of mature leaves. In phloem, a larger conductive region was observed in the wet season, when the trees were in full foliage. The secondary phloem did not exhibit any incremental zone marker; however, we found that the axial parenchyma tends to form irregular bands.     

What is disjunctive xylem parenchyma? A case study of the African tropical hardwood Okoubaka aubrevillei (Santalaceae)

The morphological variation and structure-function relationships of xylem parenchyma still remain open to discussion. We analyzed the three-dimensional structure of a poorly known type of xylem parenchyma with disjunctive walls in the tropical hardwood Okoubaka aubrevillei (Santalaceae). Disjunctive cells occurred among the apotracheal parenchyma cells and at connections between axial and ray parenchyma cells. The disjunctive cells were partly detached one from another, but their tubular structures connected them into a continuous network of axial and ray parenchyma. The connecting tubules had thick secondary walls and simple pits with plasmodesmata at the points where one cell contacted a tubule of another cell. The imperforate tracheary elements of the ground tissue were seven times longer than the axial parenchyma strands, a fact that supports a hypothesis that parenchyma cells develop disjunctive walls because they are pulled apart and partly separated during the intrusive growth of fibers. We discuss unresolved details of the formation of disjunctive cell walls and the possible biomechanical advantage of the wood with disjunctive parenchyma: the proportion of tissue that improves mechanical strength is increased by the intrusive elongation of fibers (thick-walled tracheids), whereas the symplastic continuum of the parenchyma is maintained through formation of disjunctive cells.     

Effects of wildfire on growth,transpiration and hydraulic properties of Pinus pinaster Aiton forest

In the last decades, forests have been seriously threatened by the effects of climate change. In Mediterranean regions, increasing fire occurrence and drought have a substantial impact on forests, possibly pushing them towards tipping points. Indeed, although trees could survive to fire events of medium severity in the short term, severe defoliation and environmental stressors could affect the capacity to transport water and to fix and use carbon, amplifying the possibility of a delayed post fire mortality. Therefore, it is urgent to better understand trees eco-physiological processes in response to wildfire in order to plan suitable forest management strategies. In this research an innovative continuous monitoring system of sap flow and micro-stationary climatic conditions of trees was combined with dendro-anatomical analyses and xylem cavitation vulnerability studies to assess fire impact on a Pinus pinaster forest affected by a severe fire and compared to an unburned control site. Xylem analyses showed that the forest-fire did not influence the xylem hydraulic parameters and did not increased vulnerability to embolism. However, burned trees with high defoliation rate, presented reduced assimilation capacity and carbon availability for growth, as well as a limited development of thicker xylem conduit walls compared to unburned trees. Furthermore, continuous monitoring highlighted an increase in sap flow of defoliated trees compared to the control ones during the hottest and driest months of the year indicating a higher stomatal conductance. Overall, our results suggested that although fire did not affect the xylem hydraulics properties of the vascular system, the severe crown damage caused growth reduction and transpiration dysfunction in the monitored period. Those effect could negatively affect the survival of burned trees, especially in areas subjected to hot and dry periods.     

Do xylem fibers affect vessel cavitation resistance?

Possible mechanical and hydraulic costs to increased cavitation resistance were examined among six co-occurring species of chaparral shrubs in southern California. We measured cavitation resistance (xylem pressure at 50% loss of hydraulic conductivity), seasonal low pressure potential (P(min)), xylem conductive efficiency (specific conductivity), mechanical strength of stems (modulus of elasticity and modulus of rupture), and xylem density. At the cellular level, we measured vessel and fiber wall thickness and lumen diameter, transverse fiber wall and total lumen area, and estimated vessel implosion resistance using (t/b)(h)(2), where t is the thickness of adjoining vessel walls and b is the vessel lumen diameter. Increased cavitation resistance was correlated with increased mechanical strength (r(2) = 0.74 and 0.76 for modulus of elasticity and modulus of rupture, respectively), xylem density (r(2) = 0.88), and P(min) (r(2) = 0.96). In contrast, cavitation resistance and P(min) were not correlated with decreased specific conductivity, suggesting no tradeoff between these traits. At the cellular level, increased cavitation resistance was correlated with increased (t/b)(h)(2) (r(2) = 0.95), increased transverse fiber wall area (r(2) = 0.89), and decreased fiber lumen area (r(2) = 0.76). To our knowledge, the correlation between cavitation resistance and fiber wall area has not been shown previously and suggests a mechanical role for fibers in cavitation resistance. Fiber efficacy in prevention of vessel implosion, defined as inward bending or collapse of vessels, is discussed.     

Drought resistance in early and late secondary successional species from a tropical dry forest: the interplay between xylem resistance to embolism,sapwood water storage and leaf shedding

The mechanisms of drought resistance that allow plants to successfully establish at different stages of secondary succession in tropical dry forests are not well understood. We characterized mechanisms of drought resistance in early and late‐successional species and tested whether risk of drought differs across sites at different successional stages, and whether early and late‐successional species differ in resistance to experimentally imposed soil drought. The microenvironment in early successional sites was warmer and drier than in mature forest. Nevertheless, successional groups did not differ in resistance to soil drought. Late‐successional species resisted drought through two independent mechanisms: high resistance of xylem to embolism, or reliance on high stem water storage capacity. High sapwood water reserves delayed the effects of soil drying by transiently decoupling plant and soil water status. Resistance to soil drought resulted from the interplay between variations in xylem vulnerability to embolism, reliance on sapwood water reserves and leaf area reduction, leading to a tradeoff of avoidance against tolerance of soil drought, along which successional groups were not differentiated. Overall, our data suggest that ranking species' performance under soil drought based solely on xylem resistance to embolism may be misleading, especially for species with high sapwood water storage capacity.     

Hydraulic architecture of deciduous and evergreen dry rainforest tree species from north-eastern Australia

Hydraulic conductivity and xylem anatomy were examined in stems of two evergreen species, Alphitonia excelsa (Fenzal) Benth. and Austromyrtus bidwillii (Benth.) Burret., and two drought-deciduous species, Brachychiton australis (Schott and Endl.) A. Terracc. and Cochlospermum gillivraei Benth., from a seasonally dry rainforest in north Queensland, Australia. The deciduous species possessed hydraulic architecture typical of drought-sensitive plants, i.e. low wood density, wider xylem vessels, higher maximal rates of sapwood specific hydraulic conductivity (K_s) and high vulnerability to drought-induced embolism. In contrast, the evergreen species had lower rates of K_h and leaf specific conductivity (K_L) but were less susceptible to embolism. The evergreen species experienced leaf water potentials <–4.0 MPa during the dry season, while the deciduous species shed their leaves before leaf water potentials declined below –2.0 MPa. Thus, the hydraulic architecture of the evergreens allows them to withstand the greater xylem pressure gradients required to maintain water transport to the canopy during the dry season. Our results are consistent with observations made in neotropical dry forests and demonstrate that drought-deciduous species with low wood density and high water storage capacity are likely to be more hydraulically efficient, but more vulnerable to embolism, than coexisting evergreens.     

Leaf phenology is associated with soil water availability and xylem traits in a tropical dry forest

In tropical dry forests, spatial heterogeneity in soil water availability is thought to determine interspecific differences in key components of resource use strategies, such as leaf phenology and xylem function. To understand the environmental drivers of variation in leaf phenology and xylem function, we explored the relation of soil water potential to topographic metrics derived from a digital elevation model. Subsequently, we compared nine xylem hydraulic, mechanical and storage traits in 18 species in three phenological classes (readily deciduous, tardily deciduous, and evergreen) in the dry tropical forest of Chamela, Mexico. Soil water potential was negatively correlated with elevation, insolation and water flow accumulation. Evergreen species characterized low-elevation moist sites, whereas deciduous species dominated hills and dry sites. Overall, evergreen species had lower xylem specific conductivity than deciduous species, and tardily deciduous species were different from readily deciduous and evergreen species in five of eight xylem traits. In dry tropical forests, water availability promotes divergence in leaf phenology and xylem traits, ranging from low wood density, evergreen species in moist sites to a combination of low wood density, readily deciduous species plus high wood density, tardily deciduous species in dry sites.     

云南哀牢山6种常绿阔叶树木质部解剖特征的轴向和径向变化

West、Brown和Enquist提出的树木水分传导的分形网络模型(简称WBE模型)认为,树木连续分枝之间的导管或管胞直径按照一定的比率均匀变细,其总的水力阻力与水分传导的路径长度无关,从而使不同部位叶片获得基本相当的水分供应。该模型对树木高生长的水力限制假说提出了置疑。为了验证WBE模型中树木导管或管胞均匀变细的假说,该文研究了云南哀牢山中湿性常绿阔叶林中6种常绿阔叶树, 腾冲栲(Castanopsis wattii)、景东石砾(Lithocarpus chintungensis)、木果石砾(L. xylocarpus)、长尾青冈(Cyclobalanopsis stewardiana)、滇木荷(Schima noronhae)和舟柄茶(Hartia sinensis)木质部解剖特征随树高和年龄的变化。对这6个树种共14株样木进行了不同高度树干圆盘和边材生长轮取样,样木的高度为15~25 m,按照常规木材解剖的处理和分析方法,在显微镜下测定木材切片的导管直径和密度等特征。结果表明:在14株样木中,有4株树木导管直径随树木高度增加呈线性减小, 1株没有明显变化,其它9株树木导管直径在树冠以下的树干部分变化幅度较小或没有明显变化,而从树冠基部往上直到树木顶端导管直径显著减小。同一植株随着高度的增加,导管密度增加并且在树冠内增加更显著。有三分之一的树木导管占边材面积的比例随树高增加没有明显变化,其余树木导管占边材面积比在树冠以上有所减小。多数树木理论比导率在树冠以下没有明显变化而在树冠基部往上显著降低。在从髓芯开始往外的20~40个年轮范围内导管直径增加显著,但大部分植株导管直径在40个年轮后趋于稳定。不同高度圆盘导管直径随形成层发育时间的变化呈相似的趋势,并且相同发育年龄的导管直径没有明显差异。该文的研究结果说明,导管直径的轴向和径向变化一定程度上补偿了水分运输阻力随树木个体增大而增加的缺陷,但是6种常绿阔叶树树干的导管基本不按一定比率均匀变细,不支持WBE模型。     

Replicated throughfall exclusion experiment in an Indonesian perhumid rainforest: wood production,litter fall and fine root growth under simulated drought

Climate change scenarios predict increases in the frequency and duration of ENSO‐related droughts for parts of South‐East Asia until the end of this century exposing the remaining rainforests to increasing drought risk. A pan‐tropical review of recorded drought‐related tree mortalities in more than 100 monitoring plots before, during and after drought events suggested a higher drought‐vulnerability of trees in South‐East Asian than in Amazonian forests. Here, we present the results of a replicated (n = 3 plots) throughfall exclusion experiment in a perhumid tropical rainforest in Sulawesi, Indonesia. In this first large‐scale roof experiment outside semihumid eastern Amazonia, 60% of the throughfall was displaced during the first 8 months and 80% during the subsequent 17 months, exposing the forest to severe soil desiccation for about 17 months. In the experiment's second year, wood production decreased on average by 40% with largely different responses of the tree families (ranging from −100 to +100% change). Most sensitive were trees with high radial growth rates under moist conditions. In contrast, tree height was only a secondary factor and wood specific gravity had no influence on growth sensitivity. Fine root biomass was reduced by 35% after 25 months of soil desiccation while fine root necromass increased by 250% indicating elevated fine root mortality. Cumulative aboveground litter production was not significantly reduced in this period. The trees from this Indonesian perhumid rainforest revealed similar responses of wood and litter production and root dynamics as those in two semihumid Amazonian forests subjected to experimental drought. We conclude that trees from paleo‐ or neotropical forests growing in semihumid or perhumid climates may not differ systematically in their growth sensitivity and vitality under sublethal drought stress. Drought vulnerability may depend more on stem cambial activity in moist periods than on tree height or wood specific gravity.     

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.