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Tension wood is a specialized xylem tissue associated with gravitropism in angiosperm trees. However, few regulators of tension wood formation have been identified. The molecular mechanisms underpinning tension wood formation remain elusive. Here, we report that a Populus KNOTTED-like homeobox gene, PagKNAT2/6b, is involved in tension wood formation and gravity response. Transgenic poplar plants overexpressing PagKNAT2/6b displayed more sensitive gravitropism than controls, as indicated by increased stem curvature. Microscopic examination revealed greater abundance of fibre cells with a gelatinous cell wall layer (G-layer) and asymmetric growth of secondary xylem in PagKNAT2/6b overexpression lines. Conversely, PagKNAT2/6b dominant repression plants exhibited decreased tension wood formation and reduced response to gravity stimulation. Moreover, sensitivity to gravity stimulation showed a negative relationship with development stage. Expression of genes related to growth and senescence was affected in PagKNAT2/6b transgenic plants. More importantly, transcription activation and electrophoretic mobility shift assays suggested that PagKNAT2/6b promotes the expression of cytokinin metabolism genes. Consistently, cytokinin content was increased in PagKNAT2/6b overexpression plants. Therefore, PagKNAT2/6b is involved in gravitropism and tension wood formation, likely via modulation of cytokinin metabolism., (© The Author(s) 2024. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Background: Drought periods are major evolutionary triggers of wood anatomical adaptive variation in Lower Tropical Montane Cloud Forests tree species. We tested the influence of historical drought events on the effects of ecological stress memory on latewood width and xylem vessel traits in two relict hickory species (Carya palmeri and Carya myristiciformis) from central-eastern Mexico. We hypothesized that latewood width would decrease during historical drought years, establishing correlations between growth and water stress conditions, and that moisture deficit during past tree growth between successive drought events, would impact on wood anatomical features. We analyzed latewood anatomical traits that developed during historical drought and pre- and post-drought years in both species., Results: We found that repeated periods of hydric stress left climatic signatures for annual latewood growth and xylem vessel traits that are essential for hydric adaptation in tropical montane hickory species., Conclusions: Our results demonstrate the existence of cause‒effect relationships in wood anatomical architecture and highlight the ecological stress memory linked with historical drought events. Thus, combined time-series analysis of latewood width and xylem vessel traits is a powerful tool for understanding the ecological behavior of hickory species., (© 2024. The Author(s).)

Future climatic scenarios forecast increasingly frequent droughts that will pose substantial consequences on tree mortality. In light of this, drought-tolerant eucalypts have been propagated; however, the severity of these conditions will invoke adaptive responses, impacting the commercially valuable wood properties. To determine what mechanisms govern the wood anatomical adaptive response, highly controlled drought experiments were conducted in Eucalyptus grandis W. Hill ex Maiden, with the tree physiology and transcriptome closely monitored. In response to water deficit, E. grandis displays an isohydric stomatal response to conserve water and enable stem growth to continue, albeit at a reduced rate. Maintaining gaseous exchange is likely a critical short-term response that drives the formation of hydraulically safer xylem. For instance, the development of significantly smaller fibers and vessels was found to increase cellular density, thereby promoting drought tolerance through improved functional redundancy, as well as implosion and cavitation resistance. The transcriptome was explored to identify the molecular mechanisms responsible for controlling xylem cell size during prolonged water deficit. Downregulation of genes associated with cell wall remodeling and the biosynthesis of cellulose, hemicellulose and pectin appeared to coincide with a reduction in cellular enlargement during drought. Furthermore, transcript levels of NAC and MYB transcription factors, vital for cell wall component biosynthesis, were reduced, while those linked to lignification increased. The upregulation of EgCAD and various peroxidases under water deficit did not correlate with an increased lignin composition. However, with the elevated cellular density, a higher lignin content per xylem cross-sectional area was observed, potentially enhancing hydraulic safety. These results support the requirement for higher density, drought-adapted wood as a long-term adaptive response in E. grandis, which is largely influenced by the isohydric stomatal response coupled with cellular expansion-related molecular processes., (© The Author(s) 2024. Published by Oxford University Press.)

Background: The mechanisms leading to dieback and death of trees under drought remain unclear. To gain an understanding of these mechanisms, addressing major empirical gaps regarding tree structure-function relations remains essential., Scope: We give reasons to think that a central factor shaping plant form and function is selection simultaneously favouring constant leaf-specific conductance with height growth and isometric (1:1) scaling between leaf area and the volume of metabolically active sink tissues ('sapwood'). Sapwood volume-leaf area isometry implies that per-leaf area sapwood volumes become transversely narrower with height growth; we call this 'stretching'. Stretching means that selection must favour increases in permeability above and beyond that afforded by tip-to-base conduit widening ("ultra-widening permeability"), via fewer and wider vessels or tracheids with larger pits or larger margo openings. Leaf area-metabolically active sink tissue isometry would mean that it is unlikely that larger trees die during drought because of carbon starvation due to greater sink-source relationships as compared to shorter plants. Instead, an increase in permeability is most plausibly associated with greater risk of embolism, and this seems a more probable explanation of the preferential vulnerability of larger trees to climate change-induced drought. Other implications of selection favouring constant per-leaf area sapwood construction and maintenance costs are departure from the da Vinci rule expectation of similar sapwood areas across branching orders, and that extensive conduit furcation in the stem seems unlikely., Conclusions: Because all these considerations impact the likelihood of vulnerability to hydraulic failure versus carbon starvation, both implicated as key suspects in forest mortality, we suggest that these predictions represent essential priorities for empirical testing., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For commercial re-use, please contact reprints@oup.com for reprints and translation rights for reprints. All other permissions can be obtained through our RightsLink service via the Permissions link on the article page on our site—for further information please contact journals.permissions@oup.com.)

Background and Aims: Understanding anatomical variations across plant phylogenies and environmental gradients is vital for comprehending plant evolution and adaptation. Previous studies on tropical woody plants have paid limited attention to quantitative differences in major xylem tissues, which serve specific roles in mechanical support (fibres), carbohydrate storage and radial conduction (radial parenchyma, rays), wood capacitance (axial parenchyma) and water transport (vessels). To address this gap, we investigate xylem fractions in 173 tropical tree species spanning 134 genera and 53 families along a 2200-m elevational gradient on Mount Cameroon, West Africa., Methods: We determined how elevation, stem height and wood density affect interspecific differences in vessel, fibre, and specific axial (AP) and radial (RP) parenchyma fractions. We focus on quantifying distinct subcategories of homogeneous or heterogeneous rays and apotracheal, paratracheal and banded axial parenchyma., Key Results: Elevation-related cooling correlated with reduced AP fractions and vessel diameters, while fibre fractions increased. Lower elevations exhibited elevated AP fractions due to abundant paratracheal and wide-banded parenchyma in tall trees from coastal and lowland forests. Vasicentric and aliform AP were predominantly associated with greater tree height and wider vessels, which might help cope with high evaporative demands via elastic wood capacitance. In contrast, montane trees featured a higher fibre proportion, scarce axial parenchyma, smaller vessel diameters and higher vessel densities. The lack of AP in montane trees was often compensated for by extended uniseriate ray sections with upright or squared ray cells or the presence of living fibres., Conclusions: Elevation gradient influenced specific xylem fractions, with lower elevations showing elevated AP due to abundant paratracheal and wide-banded parenchyma, securing greater vessel-to-parenchyma connectivity and lower embolism risk. Montane trees featured a higher fibre proportion and smaller vessel diameters, which may aid survival under greater environmental seasonality and fire risk., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For commercial re-use, please contact reprints@oup.com for reprints and translation rights for reprints. All other permissions can be obtained through our RightsLink service via the Permissions link on the article page on our site—for further information please contact journals.permissions@oup.com.)

Large trees in plantations generally produce more wood per unit of resource use than small trees. Two processes may account for this pattern: greater photosynthetic resource use efficiency or greater partitioning of carbon to wood production. We estimated gross primary production (GPP) at the individual scale by combining transpiration with photosynthetic water-use efficiency of Eucalyptus trees. Aboveground production fluxes were estimated using allometric equations and modeled respiration; total belowground carbon fluxes (TBCF) were estimated by subtracting aboveground fluxes from GPP. Partitioning was estimated by dividing component fluxes by GPP. Dominant trees produced almost three times as much wood as suppressed trees. They used 25 ± 10% (mean ± SD) of their photosynthates for wood production, whereas suppressed trees only used 12 ± 2%. By contrast, dominant trees used 27 ± 19% of their photosynthate belowground, whereas suppressed trees used 58 ± 5%. Intermediate trees lay between these extremes. Photosynthetic water-use efficiency of dominant trees was c. 13% greater than the efficiency of suppressed trees. Suppressed trees used more than twice as much of their photosynthate belowground and less than half as much aboveground compared with dominant trees. Differences in carbon partitioning were much greater than differences in GPP or photosynthetic water-use efficiency., (© 2024 The Authors. New Phytologist © 2024 New Phytologist Foundation.)

Understanding the capacity of temperate trees to acclimate to limited soil water has become essential in the face of increasing drought risk due to climate change. We documented seasonal - or phenological - patterns in acclimation to water deficit stress in stems and leaves of tree species spanning the angiosperm phylogeny. Over 3 yr of field observations carried out in two US arboreta, we measured stem vulnerability to embolism (36 individuals of 7 Species) and turgor loss point (119 individuals of 27 species) over the growing season. We also conducted a growth chamber experiment on 20 individuals of one species to assess the mechanistic relationship between soil water restriction and acclimation. In three-quarters of species measured, plants became less vulnerable to embolism and/or loss of turgor over the growing season. We were able to stimulate this acclimatory effect by withholding water in the growth chamber experiment. Temperate angiosperms are capable of acclimation to soil water deficit stress, showing maximum vulnerability to soil water deficits following budbreak and becoming more resilient to damage over the course of the growing season or in response to simulated drought. The species-specific tempo and extent of this acclimatory potential constitutes preadaptive climate change resilience., (© 2024 The Authors. New Phytologist © 2024 New Phytologist Foundation.)

Understanding and predicting recruitment in species-rich plant communities requires identifying functional determinants of both density-independent performance and interactions. In a common-garden field experiment with 25 species of the woody plant genus Protea, we varied the initial spatial and taxonomic arrangement of seedlings and followed their survival and growth during recruitment. Neighbourhood models quantified how six key functional traits affect density-independent performance, interaction effects and responses. Trait-based neighbourhood models accurately predicted individual survival and growth from the initial spatial and functional composition of species-rich experimental communities. Functional variation among species caused substantial variation in density-independent survival and growth that was not correlated with interaction effects and responses. Interactions were spatially restricted but had important, predominantly competitive, effects on recruitment. Traits increasing the acquisition of limiting resources (water for survival and soil P for growth) mediated trade-offs between interaction effects and responses. Moreover, resprouting species had higher survival but reduced growth, likely reinforcing the survival-growth trade-off in adult plants. Resource acquisition of juvenile plants shapes Protea community dynamics with acquisitive species with strong competitive effects suffering more from competition. Together with functional determinants of density-independent performance, this makes recruitment remarkably predictable, which is critical for efficient restoration and near-term ecological forecasts of species-rich communities., (© 2023 The Authors. New Phytologist © 2023 New Phytologist Foundation.)

The predicted increase of drought intensity in South-East Asia has raised concern about the sustainability of rubber (Hevea brasiliensis Müll. Arg.) cultivation. In order to quantify the degree of phenotypic plasticity in this important tree crop species, we analysed a set of wood and leaf traits related to the hydraulic safety and efficiency in PB260 clones from eight small-holder plantations in Jambi province, Indonesia, representing a gradient in local microclimatic and edaphic conditions. Across plots, branch embolism resistance (P50) ranged from -2.14 to -2.58 MPa. The P50 and P88 values declined, and the hydraulic safety margin increased, with an increase in the mean annual vapour pressure deficit (VPD). Among leaf traits, only the changes in specific leaf area were related to the differences in evaporative demand. These variations of hydraulic trait values were not related to soil moisture levels. We did not find a trade-off between hydraulic safety and efficiency, but vessel density (VD) emerged as a major trait associated with both safety and efficiency. The VD, and not vessel diameter, was closely related to P50 and P88 as well as to specific hydraulic conductivity, the lumen-to-sapwood area ratio and the vessel grouping index. In conclusion, our results demonstrate some degree of phenotypic plasticity in wood traits related to hydraulic safety in this tropical tree species, but this is only in response to the local changes in evaporative demand and not soil moisture. Given that VPD may increasingly limit plant growth in a warmer world, our results provide evidence of hydraulic trait changes in response to a rising evaporative demand., (© The Author(s) 2023. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Evolutionary radiations of woody taxa within arid environments were made possible by multiple trait innovations including deep roots and embolism-resistant xylem, but little is known about how these traits have coevolved across the phylogeny of woody plants or how they jointly influence the distribution of species. We synthesized global trait and vegetation plot datasets to examine how rooting depth and xylem vulnerability across 188 woody plant species interact with aridity, precipitation seasonality, and water table depth to influence species occurrence probabilities across all biomes. Xylem resistance to embolism and rooting depth are independent woody plant traits that do not exhibit an interspecific trade-off. Resistant xylem and deep roots increase occurrence probabilities in arid, seasonal climates over deep water tables. Resistant xylem and shallow roots increase occurrence probabilities in arid, nonseasonal climates over deep water tables. Vulnerable xylem and deep roots increase occurrence probabilities in arid, nonseasonal climates over shallow water tables. Lastly, vulnerable xylem and shallow roots increase occurrence probabilities in humid climates. Each combination of trait values optimizes occurrence probabilities in unique environmental conditions. Responses of deeply rooted vegetation may be buffered if evaporative demand changes faster than water table depth under climate change., (© 2023 The Authors. New Phytologist © 2023 New Phytologist Foundation.)

Wood performs several functions to ensure tree survival and carbon allocation to a finite stem volume leads to trade-offs among cell types. It is not known to what extent these trade-offs modify functional trade-offs and if they are consistent across climates and evolutionary lineages. Twelve wood traits were measured in stems and coarse roots across 60 adult angiosperm tree species from temperate, Mediterranean and tropical climates. Regardless of climate, clear trade-offs occurred among cellular fractions, but did not translate into specific functional trade-offs. Wood density was negatively related to hydraulic conductivity (K th ) in stems and roots, but was not linked to nonstructural carbohydrates (NSC), implying a functional trade-off between mechanical integrity and transport but not with storage. NSC storage capacity was positively associated with K th in stems and negatively in roots, reflecting a potential role for NSC in the maintenance of hydraulic integrity in stems but not in roots. Results of phylogenetic analyses suggest that evolutionary histories cannot explain covariations among traits. Trade-offs occur among cellular fractions, without necessarily modifying trade-offs in function. However, functional trade-offs are driven by coordinated changes among xylem cell types depending on the dominant role of each cell type in stems and roots., (© 2023 The Authors. New Phytologist © 2023 New Phytologist Foundation.)

Molecular studies indicate that Penaeaceae, Oliniaceae, and the monospecific families Alzateaceae and Rhynchocalycaceae form a clade of Myrtales. Of these four families, Penaeaceae have tracheids with vestured pits, whereas the others have septate fibers lacking vestures; all have vestured pits in vessels. Tracheid presence in Penaeaceae may be related to the arid South African habitats of the family. Presence of vestures on tracheids in families with vestured vessel pits is one indication that imperforate elements are tracheids and are conductive cells, whereas fiber-tracheids and libriform fibers are non-conductive. Tracheids occur widely in angiosperms and may be plesiomorphies or apomorphies. Combretaceae, the first branch of the Myrtales clade, has a great diversity of vesture features in vessels compared to the Penaeaceae alliance families. Alzatea has vestures that spread over the inside of the vessels, whereas in most taxa of the alliance, vestures are confined to the pit cavities and pit apertures. Vestures in the alliance tend to be globular in shape, and are bridged together by strands of wall material. Lignotubers and roots in Penaeaceae have vestures much like those in stems. Only a few species and genera (notably Alzatea) of the alliance have vesture features the pattern of which correlates with the current taxonomic system. Vestured pits should be viewed from the inside surface of vessels as well as the outer surface, and although sectional views of vestured pits are infrequent, they are very informative. Studies that explore diversity from one order or family to another are needed and offer opportunities for understanding the evolutionary significance of this feature. [ABSTRACT FROM AUTHOR]

The nature of conduction involves movement of a liquid (under tension or pressure) through a solid (cell walls necessary to direct the liquid and provide mechanical strength). The numerous consequences of the liquid/solid nature of the conductive interface in plants can be viewed as a series of conflicting requirements that are resolved by various mechanisms. For example, the types of mechanical strength conferred by thicker cell walls (latewood) run counter to optimal conduction (earlywood). Conflict resolution situations are examined with light microscopy and SEM to show in detail not merely conflicting requirements but the various types of resolution in various conifers. Abies is presented as exemplary of a cool temperate conifer with numerous aspects to earlywood/latewood structure. Tropical conifers (Araucaria) present different compromises; the riparian conifer Dacrydium guillauminii has only earlywood; the parasitic conifer Parasitaxus has only latewood. Particular conifers have only some of the features by which latewood differs from earlywood. Cell dimorphism is only one aspect of resolution of conflicting requirements; others include modifications in pit size, shape, and density; the nature of the pit membrane; the nature of the pit cavity, pit border and pit aperture; and surface relief (warty layer) of the tracheid wall. The invention of coniferous bordered pits involves a circular shape, so that tension on the margo strands is equal, and thus the pit can be closed. These factors and margo pore maximization necessitate expending a large amount of space to pits in earlywood, the strength of which is thereby lessened and must be compensated by greater wall strength in latewood. The paper concludes with a series of twenty features which represent resolutions of conflicting requirements in terms of anatomical structure. Wood physiological literature is integrated with the anatomical observations. [ABSTRACT FROM AUTHOR]

Bush encroachment affects much of the Namibian woodland landscape, causing significant loss of open savannah habitat and farm profits. Thinning of the trees/shrubs is recommended; however, research is required to identify the overall efficacy and effects of this method on the woodland habitat. We aimed to examine the effect of the thinning strategy applied on the vegetation structure of encroaching tree/shrub species, as well as the sighting lines of the habitat. Vegetation surveys were done on three freehold farms in north-central Namibia. The study utilised a combination of a blocked and split-plot study design: each block consisted of a pair of thinned and non-thinned plots with multiple subplots. Thinned plots had been manually thinned, with a post-thinning age of three years or more. Results revealed that tree/shrub abundance differed between species; thinned areas had the least abundance and overall species-treatment interactions were significant. Thinning caused a significant reduction in overall tree/shrub densities, settling within the recommended range for the area. Thinning also significantly reduced the average tree/shrub height, canopy area, medium-sized trees/shrubs, and increased sighting lines. This confirms a bush encroachment mitigation strategy that favours grass cover, and wildlife that rely on longer sighting lines for safety or when hunting., (© 2023. The Author(s).)

Accidental (strimmer, mower) or malicious girdling of ornamental trees is a frequent and underestimated problem in urban landscapes. No guidelines exist for landscape managers on the likelihood of fatal damage according to time of girdle, i.e., season (spring, winter) and extent of girdling. This report investigated the effect of stem girdling (0%, 25%, 50%, 75%, 100%) performed either during winter or spring on semi-mature birch ( Betula pendula Roth.) and English oak ( Quercus robor L) trees over a five-year period. 1. All English oak survived any amount of girdling irrespective if girdled in spring or winter. Silver birch was more sensitive to girdling with 100% and 40% mortality recorded after a 100% winter and spring girdle, respectively. 2. Tree vitality (leaf chlorophyll fluorescence, leaf chlorophyll content, crown leaf yellowing, nitrogen content) and growth (girth increment, die-back, number of epicormics above and below the girdle) of both tree species was not detrimentally affected following a 25% and 50% girdle. 3. Epicormic production was more pronounced in English oaks compared to silver birch. 4. Spring girdled oak typically had lower tree vitality and growth values than winter girdled trees indicating an influence of time of girdling. Such a response was not recorded in silver birch where time of girdling induced similar effects on tree growth and vitality. 5. Growth and vitality measurements of surviving trees of both species were, in virtually all cases, statistically comparable with non-girdled trees by the end of the third growing season. Such a result indicates girdling >50% influences growth and vitality of both English oak and silver birch for three years. [ABSTRACT FROM AUTHOR]

The diversity of expression in axial parenchyma (or lack of it) in woods is reviewed and synthesized with recent work in wood physiology, and questions and hypotheses relative to axial parenchyma anatomy are offered. Cell shape, location, abundance, size, wall characteristics and contents are all characteristics for the assessment of the physiological functions of axial parenchyma, a tissue that has been neglected in the consideration of how wood histology has evolved. Axial parenchyma occurrence should be considered with respect to mechanisms for the prevention and reversal of embolisms in tracheary elements. This mechanism complements cohesion-tension-based water movement and root pressure as a way of maintaining flow in xylem. Septate fibres can substitute for axial parenchyma ('axial parenchyma absent') and account for water movement in xylem and for the supply of carbohydrate abundance underlying massive and sudden events of foliation, flowering and fruiting, as can fibre dimorphism and the co-occurrence of septate fibres and axial parenchyma. Rayless woods may or may not contain axial parenchyma and are informative when analysing parenchyma function. Interconnections between ray and axial parenchyma are common, and so axial and radial parenchyma must be considered as complementary parts of a network, with distinctive but interactive functions. Upright ray cells and more numerous rays per millimetre enhance interconnection and are more often found in woods that contain tracheids. Vesselless woods in both gymnosperms and angiosperms have axial parenchyma, the distribution of which suggests a function in osmotic water shifting. Water and photosynthate storage in axial parenchyma may be associated with seasonal changes and with succulent or subsucculent modes of construction. Apotracheal axial parenchyma distribution often demonstrates storage functions that can be read independently of osmotic water shifting capabilities. Axial parenchyma may serve to both enhance mechanical strength or, when parenchyma is thin-walled, as a tissue that adapts to volume change with a change in water content. Other functions of axial parenchyma (contributing resistance to pathogens; a site for the recovery of physical damage) are considered. The diagnostic features of axial parenchyma and septate fibres are reviewed in order to clarify distinctions and to aid in cell type identification. Systematic listings are given for particular axial parenchyma conditions (e.g. axial parenchyma 'absent' with septate fibres substituting). A knowledge of the axial parenchyma information presented here is desirable for a full understanding of xylem function. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 177, 291-321. [ABSTRACT FROM AUTHOR]

Whether sources or sinks control wood growth remains debated with a paucity of evidence from mature trees in natural settings. Here, we altered carbon supply rate in stems of mature red maples (Acer rubrum) within the growing season by restricting phloem transport using stem chilling; thereby increasing carbon supply above and decreasing carbon supply below the restrictions, respectively. Chilling successfully altered nonstructural carbon (NSC) concentrations in the phloem without detectable repercussions on bulk NSC in stems and roots. Ring width responded strongly to local variations in carbon supply with up to seven-fold differences along the stem of chilled trees; however, concurrent changes in the structural carbon were inconclusive at high carbon supply due to large local variability of wood growth. Above chilling-induced bottlenecks, we also observed higher leaf NSC concentrations, reduced photosynthetic capacity, and earlier leaf coloration and fall. Our results indicate that the cambial sink is affected by carbon supply, but within-tree feedbacks can downregulate source activity, when carbon supply exceeds demand. Such feedbacks have only been hypothesized in mature trees. Consequently, these findings constitute an important advance in understanding source-sink dynamics, suggesting that mature red maples operate close to both source- and sink-limitation in the early growing season., (© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.)

Background and Aims: Xylem is a crucial tissue for plant survival, performing the functions of water transport, mechanical support and storage. Functional trade-offs are a result of the different assemblages of xylem cell types within a certain wood volume. We assessed how the volume allocated to different xylem cell types can be associated with wood functional trade-offs (hydraulics, mechanical and storage) in species from the Cerrado, the Brazilian savanna. We also assessed the xylem anatomical characters linked to wood density across species., Methods: We analysed cross-sections of branches collected from 75 woody species belonging to 42 angiosperm families from the Cerrado. We estimated the wood volume fraction allocated to different cell types and performed measurements of vessel diameter and wood density., Key Results: The largest volume of wood is allocated to fibres (0.47), followed by parenchyma (0.33) and vessels (0.20). Wood density is positively correlated to cell wall (fibre and vessel wall), and negatively to the fractions of fibre lumen and gelatinous fibres. We observed a trade-off between hydraulics (vessel diameter) and mechanics (cell wall fraction), and between mechanics and storage (parenchyma fraction). The expected positive functional relationships between hydraulics (vessel diameter) and water and carbohydrate storage (parenchyma and fibre lumen fractions) were not detected, though larger vessels are linked to a larger wood volume allocated to gelatinous fibres., Conclusions: Woody species from the Cerrado show evidence of functional trade-offs between water transport, mechanical support and storage. Gelatinous fibres might be potentially linked to water storage and release by their positive relationship to increased vessel diameter, thus replacing the functional role of parenchyma and fibre lumen cells. Species can profit from the increased mechanical strength under tension provided by the presence of gelatinous fibres, avoiding expensive investments in high wood density., (© The Author(s) 2022. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

There is a long-standing idea that the timing of leaf production in seasonally cold climates is linked to xylem anatomy, specifically vessel diameter because of the hydraulic requirements of expanding leaves. We tested for a relationship between the timing of leaf out and vessel diameter in 220 plants in three common gardens accounting for species' phylogenetic relationships. We investigated how vessel diameter related to wood porosity, plant height and leaf length. We also used dye perfusion tests to determine whether plants relied on xylem produced during the previous growing season at the time of leaf out. In all three gardens, there was later leaf out in species with wider vessels. Ring-porous species had the widest vessels, exhibited latest leaf out and relied less on xylem made during the previous growing season than diffuse-porous species. Wood anatomy and leaf phenology did not exhibit a phylogenetic signal. The timing of leaf out is correlated with wood anatomy across species regardless of species' geographic origin and phylogenetic relationships. This correlation could be a result of developmental and physiological links between leaves and wood or tied to a larger safety efficiency trade-off., (© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.)

The increased frequency and intensity of drought pose great threats to the survival of trees, especially in dioecious tree species with sexual differences in mortality and biased sex ratios. The sex-specific mechanisms underlying stem xylem anatomy and function and carbon metabolism in drought resistance and recovery were investigated in dioecious Populus cathayana Rehder. The sex-specific drought resistance and subsequent recovery were linked to the xylem anatomy and carbon metabolism. Females had a greater xylem vessel area per vessel, biomass and theoretically hydraulic efficiency under well-watered conditions. Conversely, males had a lower xylem lumen area, but greater vessel numbers, and a higher cell wall thickness, suggesting a theoretically conservative water-use strategy and drought resistance. The recovery of photosynthetic ability after drought in males was largely dependent on the recovery of xylem function and the regulation of the xylem carbohydrate metabolism. Additionally, the number of upregulated genes related to xylem cell wall biogenesis was greater in males relative to females under drought stress and subsequent rewatering, which facilitated drought resistance and xylem function restoration in males. These results suggested that sex-specific drought resistance and restoration were related to xylem anatomy and function, carbohydrate metabolism and cell turgor maintenance., (© The Author(s) 2022. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Due to massive energetic investments in woody support structures, trees are subject to unique physiological, mechanical, and ecological pressures not experienced by herbaceous plants. Despite a wealth of studies exploring trait relationships across the entire plant kingdom, the dominant traits underpinning these unique aspects of tree form and function remain unclear. Here, by considering 18 functional traits, encompassing leaf, seed, bark, wood, crown, and root characteristics, we quantify the multidimensional relationships in tree trait expression. We find that nearly half of trait variation is captured by two axes: one reflecting leaf economics, the other reflecting tree size and competition for light. Yet these orthogonal axes reveal strong environmental convergence, exhibiting correlated responses to temperature, moisture, and elevation. By subsequently exploring multidimensional trait relationships, we show that the full dimensionality of trait space is captured by eight distinct clusters, each reflecting a unique aspect of tree form and function. Collectively, this work identifies a core set of traits needed to quantify global patterns in functional biodiversity, and it contributes to our fundamental understanding of the functioning of forests worldwide., (© 2022. The Author(s).)

Tree species display a wide variety of water-use strategies, growth rates and capacity to tolerate drought. However, if we want to forecast species capacity to cope with increasing aridity and drought, we need to identify which measurable traits confer resilience to drought across species. Here, we use a global tree ring network (65 species; 1931 site series of ring-width indices-RWI) to evaluate the relationship of long-term growth-drought sensitivity (RWI-SPEI drought index relationship) and short-term growth response to extreme drought episodes (resistance, recovery and resilience indices) with functional traits related to leaf, wood and hydraulic properties. Furthermore, we assess the influence of climate (temperature, precipitation and climatic water deficit) on these trait-growth relationships. We found a close correspondence between the long-term relationship between RWI and SPEI and resistance and recovery of tree growth to severe drought episodes. Species displaying a stronger RWI-SPEI relationship to drought and low resistance and high recovery to extreme drought episodes tended to have a higher wood density (WD) and more negative leaf minimum water potential (Ψmin). Such associations were largely maintained when accounting for direct climate effects. Our results indicate that, at a cross-species level and global scale, wood and hydraulic functional traits explain species' growth responses to drought at short- and long-term scales. These trait-growth response relationships can improve our understanding of the cross-species capacity to withstand climate change and inform models to better predict drought effects on forest ecosystem dynamics., (© 2022 John Wiley & Sons Ltd.)

Recent advances in wood physiology, molecular phylogeny, and ultrastructure (chiefly scanning electron microscopy, SEM), as well as important new knowledge in traditional fields, provide the basis for a new vision of how wood evolves. Woody angiosperms have, in the main, shifted from conductive safety to conductive efficiency (with many variations and modifications) and from ability to resist cavitation (low vulnerability) to ability to refill vessels. The invention of the vessel was a kind of dimorphism (vessel elements plus tracheids) that permitted division of labor and many kinds of wood repatterning that suit conductive safety-efficiency trade-offs. Angiosperms were primarily adapted to mesic habitats but were not failures or 'unstable.' They have survived to the present in such habitats well, along with older structural adaptations (e.g., the scalariform perforation plate) that are still suited to such habitats. These 'primitive' features are evident in earlier branchings of phylogenetic trees based on multiple genes. Older features may still be functional and thus persist, although newer formulations are overriding in effect. There are, however, numerous instances of 'breakouts' in a number of clades (ecological iterations and bursts of speciation and diversification related to new ways of dealing with water economy), whereas in other branchings, other clades show ecological stasis over long periods of time. Newer physiological and anatomical mechanisms have permitted entry into habitats with marked fluctuation in moisture availability. Wood evolves progressively, and literal character state reversal may be unusual: genomic and developmental information holds answers to these changes. Wood is a complex tissue, and each of the histological components shows polymorphism as an evolutionary mechanism. Cell types within wood evolve collaboratively. Shifts in wood features (e.g., simplification of the scalariform perforation plate) are commonly homoplastic. Manifold changes in habit and in leaf physiology, morphology, and anatomy accompany wood evolution, and wood should be studied with relationship to real-world ecology, information that cannot be gleaned from literature or other secondary sources. Heterochrony (protracted juvenilism, accelerated adulthood) characterizes angiosperm xylem extensively, far more so than in other vascular plants, and these mechanisms have resulted in many remarkable changes (e.g., monocots have permanently juvenile xylem, woody trees represent accelerated adulthood). Understanding the many successful features of angiosperm wood evolution must ultimately rest on syntheses. [ABSTRACT FROM AUTHOR]

Parenchyma is an important component of the secondary xylem. It has multiple functions and its fraction is known to vary substantially across angiosperm species. However, the physiological significance of this variation is not yet fully understood. Here, we examined how different types of parenchyma (ray parenchyma [RP], axial parenchyma [AP] and AP in direct contact with vessels [APV]) are coordinated with three essential xylem functions: water conduction, storage of non-structural carbohydrate (NSC) and mechanical support. Using branch sapwood of 15 co-occurring drought-adapted woody species from the subtropical Bonin Islands, Japan, we quantified 10 xylem anatomical traits and examined their linkages to hydraulic properties, storage of soluble sugars and starch and sapwood density. The fractions of APV and AP in the xylem transverse sections were positively correlated with the percentage loss of conductivity in the native condition, whereas that of RP was negatively correlated with the maximum conductivity across species. Axial and ray parenchyma fractions were positively associated with concentrations of starch and NSC. The fraction of parenchyma was independent of sapwood density, regardless of parenchyma type. We also identified a negative relationship between hydraulic conductivity and NSC storage and sapwood density, mirroring the negative relationship between the fractions of parenchyma and vessels. These results suggest that parenchyma fraction underlies species variation in xylem hydraulic and carbon use strategies, wherein xylem with a high fraction of AP may adopt an embolism repair strategy through an increased starch storage with low cavitation resistance., (© The Author(s) 2021. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Despite extensive knowledge about vessel element growth and the determination of the axial course of vessels, these processes are still not fully understood. They are usually explained as resulting primarily from hormonal regulation in stems. This review focuses on an increasingly discussed aspect - mechanical conditions in the vascular cambium. Mechanical conditions in cambial tissue are important for the growth of vessel elements, as well as other cambial derivatives. In relation to the type of stress acting on cambial cells (compressive versus tensile stress) we: (i) discuss the shape of the enlarging vessel elements observed in anatomical sections; (ii) present hypotheses regarding the location of intrusive growth of vessel elements and cambial initials; (iii) explain the relationship between the growth of vessel elements and fibres; and (iv) consider the effect of mechanical stress in determining the course of a vessel. We also highlight the relationship between mechanical stress and transport of the most extensively studied plant hormone - auxin. We conclude that the integration of a biomechanical factor with the commonly acknowledged hormonal regulation could significantly enhance the analysis of the formation of vessel elements as well as entire vessels, which transport water and minerals in numerous plant species., (© 2021 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society.)

Fungal pathogens have evolved combinations of plant cell-wall-degrading enzymes (PCWDEs) to deconstruct host plant cell walls (PCWs). An understanding of this process is hoped to create a basis for improving plant biomass conversion efficiency into sustainable biofuels and bioproducts. Here, an approach integrating enzyme activity assay, biomass pretreatment, field emission scanning electron microscopy (FESEM), and genomic analysis of PCWDEs were applied to examine digestibility or degradability of selected woody and herbaceous biomass by pathogenic fungi. Preferred hydrolysis of apple tree branch, rapeseed straw, or wheat straw were observed by the apple-tree-specific pathogen Valsa mali , the rapeseed pathogen Sclerotinia sclerotiorum , and the wheat pathogen Rhizoctonia cerealis , respectively. Delignification by peracetic acid (PAA) pretreatment increased PCW digestibility, and the increase was generally more profound with non-host than host PCW substrates. Hemicellulase pretreatment slightly reduced or had no effect on hemicellulose content in the PCW substrates tested; however, the pretreatment significantly changed hydrolytic preferences of the selected pathogens, indicating a role of hemicellulose branching in PCW digestibility. Cellulose organization appears to also impact digestibility of host PCWs, as reflected by differences in cellulose microfibril organization in woody and herbaceous PCWs and variation in cellulose-binding domain organization in cellulases of pathogenic fungi, which is known to influence enzyme access to cellulose. Taken together, this study highlighted the importance of chemical structure of both hemicelluloses and cellulose in host PCW digestibility by fungal pathogens.

Xylem tracheids are the channels for water transport in conifer. Tracheid flow resistance is composed of tracheid lumen resistance and pit resistance. The single tracheid structure parameters in the stem and root of Sabina chinensis were obtained by dissociation and slicing, combined with numerical simulation to analyze the tracheid flow resistance characteristics. The results showed that the tracheid lumen resistance was determined by the tracheid width and tracheid length. The pit resistance was determined by the number of pits and single pit resistance. The single pit resistance was composed of four elements: the secondary cell wall, the border, the margo and the torus. The margo contributed a relatively large fraction of flow resistance, while the torus, the border and the secondary cell wall formed a small fraction. The size and position of the pores in the margo had a significant effect on the fluid velocity. The number of pits were proportional to tracheid length. The power curve, S-curve and inverse curve were fitted the scatter plot of total pit resistance, total resistance, total resistivity, which was found that there were the negative correlation between them. The three scatter plot values were larger in the stem than in the root, indicating that the tracheid structure in the root was more conducive to water transport than the stem. The ratio of tracheid lumen resistance to pit resistance mainly was less than 0.6 in the stem and less than 1 in the root, indicating that the pit resistance was dominant in the total resistance of the stem and root., Competing Interests: no.

Cadmium (Cd 2+ ) pollution occurring in salt-affected soils has become an increasing environmental concern in the world. Fast-growing poplars have been widely utilized for phytoremediation of soil contaminating heavy metals (HMs). However, the woody Cd 2+ -hyperaccumulator, Populus × canescens , is relatively salt-sensitive and therefore cannot be directly used to remediate HMs from salt-affected soils. The aim of the present study was to testify whether colonization of P. × canescens with ectomycorrhizal (EM) fungi, a strategy known to enhance salt tolerance, provides an opportunity for affordable remediation of Cd 2+ -polluted saline soils. Ectomycorrhization with Paxillus involutus strains facilitated Cd 2+ enrichment in P. × canescens upon CdCl 2 exposures (50 μM, 30 min to 24 h). The fungus-stimulated Cd 2+ in roots was significantly restricted by inhibitors of plasmalemma H + -ATPases and Ca 2+ -permeable channels (CaPCs), but stimulated by an activator of plasmalemma H + -ATPases. NaCl (100 mM) lowered the transient and steady-state Cd 2+ influx in roots and fungal mycelia. Noteworthy, P. involutus colonization partly reverted the salt suppression of Cd 2+ uptake in poplar roots. EM fungus colonization upregulated transcription of plasmalemma H + -ATPases ( PcHA4 , 8 , 11 ) and annexins ( Pc ANN1 , 2 , 4 ), which might mediate Cd 2+ conductance through CaPCs. EM roots retained relatively highly expressed PcHAs and PcANNs , thus facilitating Cd 2+ enrichment under co-occurring stress of cadmium and salinity. We conclude that ectomycorrhization of woody hyperaccumulator species such as poplar could improve phytoremediation of Cd 2+ in salt-affected areas.

Tension wood (TW) is a specialized xylem tissue formed in angiosperm trees under gravitational stimulus or mechanical stresses (e.g., bending). The genetic regulation that underlies this important mechanism remains poorly understood. Here, we used laser capture microdissection of stem xylem cells coupled with full transcriptome RNA-sequencing to analyze TW formation in Populus trichocarpa . After tree bending, PtrLBD39 was the most significantly induced transcription factor gene; it has a phylogenetically paired homolog, PtrLBD22 . CRISPR-based knockout of PtrLBD39/22 severely inhibited TW formation, reducing cellulose and increasing lignin content. Transcriptomic analyses of CRISPR-based PtrLBD39/22 double mutants showed that these two genes regulate a set of TW-related genes. Chromatin immunoprecipitation sequencing (ChIP-seq) was used to identify direct targets of PtrLBD39. We integrated transcriptomic analyses and ChIP-seq assays to construct a transcriptional regulatory network (TRN) mediated by PtrLBD39. In this TRN, PtrLBD39 directly regulates 26 novel TW-responsive transcription factor genes. Our work suggests that PtrLBD39 and PtrLBD22 specifically control TW formation by mediating a TW-specific TRN in Populus ., (© 2021 The Author(s).)

Veteran hollow trees are keystone structures in ecosystems and provide important habitat for a diverse set of organisms, many of which are involved in the process of decomposition. Since veteran trees are 'islands' of high biodiversity, they provide a unique system in which to study the relationship between biodiversity and decomposition of wood. We tested this relationship with a balanced experiential design, where we quantified the taxonomic and functional diversity of beetles directly involved in the process of decomposing wood, and measured the decomposition of experimentally added bundles of small diameter wood around 20 veteran trees and 20 nearby young trees in southern Norway. We found that the diversity (both taxonomic and functional) of wood-decomposing beetles was significantly higher around the veteran trees, and beetle communities around veteran trees consisted of species with a greater preference for larger diameter wood. We extracted few beetles from the experimentally added wood bundles, regardless of the tree type that they were placed near, but decomposition rates were significantly lower around veteran trees. We speculate that slower decomposition rates around veteran trees could have been a result of a greater diversity of competing fungi, which has been found to decrease decay rates. Veteran trees provide an ecological legacy within anthropogenic landscapes, enhance biodiversity and influence wood decomposition. Actions to protect veteran trees are urgently needed in order to save these valuable organisms and their associated biodiversity., Competing Interests: The authors have declared that no competing interests exist.

Defining plant hydraulic traits is central to the quantification of ecohydrological processes ranging from land-atmosphere interactions, to tree mortality and water-carbon budgets. A key plant trait is the xylem specific hydraulic conductivity (K x ), that describes the plant's vascular system capacity to transport water. While xylem's vessels and tracheids are dead upon maturity, the xylem is neither inert nor deadwood, various components of the sapwood and surrounding tissue remaining alive and functional. Moreover, the established definition of K x assumes linear relations between water flux and pressure gradient by tacitly considering the xylem as a "passive conduit". Here, we re-examine this notion of an inert xylem by systematically characterizing xylem flow in several woody plants using K x measurements under constant and cyclic pressure gradients. Results show a temporal and pressure gradient dependence of K x . Additionally, microscopic features in "living branches" are irreversibly modified upon drying of the xylem, thus differentiating the macroscopic definition of K x for living and dead xylem. The findings highlight the picture of the xylem as a complex and delicate conductive system whose hydraulic behaviour transcends a passive gradient-based flow. The study sheds new light on xylem conceptualization, conductivity measurement protocols, in situ long-distance water transport and ecosystem modelling., (© 2020 John Wiley & Sons Ltd.)

Water released from wood during transpiration (capacitance) can meaningfully affect daily water use and drought response. To provide context for better understanding of capacitance mechanisms, we investigated links between capacitance and wood anatomy. On twigs of 30 temperate angiosperm tree species, we measured day capacitance (between predawn and midday), water content, wood density, and anatomical traits, that is, vessel dimensions, tissue fractions, and vessel-tissue contact fractions (fraction of vessel circumference in contact with other tissues). Across all species, wood density (WD) and predawn lumen volumetric water content (VWC L-pd ) together were the strongest predictors of day capacitance (r 2 adj = .44). Vessel-tissue contact fractions explained an additional ~10% of the variation in day capacitance. Regression models were not improved by including tissue lumen fractions. Among diffuse-porous species, VWC L-pd and vessel-ray contact fraction together were the best predictors of day capacitance, whereas among semi/ring-porous species, VWC L-pd , WD and vessel-fibre contact fraction were the best predictors. At predawn, wood was less than fully saturated for all species (lumen relative water content = 0.52 ± 0.17). Our findings imply that day capacitance depends on the amount of stored water, tissue connectivity and the bulk wood properties arising from WD (e.g., elasticity), rather than the fraction of any particular tissue., (© 2020 John Wiley & Sons Ltd.)

Understanding the determinants of range location and size is fundamental to our understanding of spatial patterns in species richness. Here, we aimed to test the role of 'climatic stability' in determining latitudinal trends in range size and as a consequence on species richness of tropical woody plants. Using primary data from 156 (0.06 ha) plots comprising 20,400 occurrences of more than 400 species of tropical woody plants, we built a biome-wide species database that covers the entire latitudinal extent of the wet-evergreen forests of the Western Ghats (8o to 20o N), India. We consolidated this database using secondary data from other published species inventories. We then calculated the range sizes and climatic niche width of woody plants to test the predictions of the climatic stability hypothesis and examined the relationship between range position and climatic tolerance of species. Our results show a significant latitudinal gradient in species richness and turnover where local and regional species richness increase monotonically from higher latitudes to lower latitudes of the Western Ghats. We found strong support for Rapoport's Rule with an increase in range size from lower to higher latitudes; our results are consistent with the predictions of the climatic stability hypothesis, where species at higher latitudes exhibited greater tolerance to temperature and rainfall seasonality. Contrary to earlier work, our findings suggest that Rapoport's Rule and the climatic stability hypothesis can operate over regional scales, and even at lower latitudes. We suggest that latitude associated climatic seasonality through its influence on species ranges, can influence latitudinal patterns in species turnover as well as species richness., Competing Interests: The authors have declared that no competing interests exist.

The remarkable mechanical stability of wood is primarily attributed to the hierarchical fibrous arrangement of the polymeric components. While the mechanisms by which fibrous cell structure and cellulose microfibril arrangements lend stiffness and strength to wood have been intensively studied, the structural origins of the relatively high splitting fracture toughness remain unclear. This study relates cellulose microfibril arrangements to splitting fracture toughness in pine wood cell walls using in situ electron microscopy and reveals a previously unknown toughening mechanism: the specific arrangement of cellulose microfibrils in the cell wall deflects cracks from the S2 layer to the S1/S2 interface, and, once there, causes the crack to be repetitively arrested and shunted along the interface in a zig-zag path. It is suggested that this natural adaptation of wood to achieve tough interfaces and then deflect and trap cracks at them can be generalized to provide design guidelines to improve toughness of high-performance and renewable engineering materials., (© 2020 The Authors. Published by WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim.)

Functional relationships between wood density and measures of xylem hydraulic safety and efficiency are ambiguous, especially in wet tropical forests. In this meta-analysis, we move beyond wood density per se and identify relationships between xylem allocated to fibers, parenchyma, and vessels and measures of hydraulic safety and efficiency. We analyzed published data of xylem traits, hydraulic properties and measures of drought resistance from neotropical tree species retrieved from 346 sources. We found that xylem volume allocation to fiber walls increases embolism resistance, but at the expense of specific conductivity and sapwood capacitance. Xylem volume investment in fiber lumen increases capacitance, while investment in axial parenchyma is associated with higher specific conductivity. Dominant tree taxa from wet forests prioritize xylem allocation to axial parenchyma at the expense of fiber walls, resulting in a low embolism resistance for a given wood density and a high vulnerability to drought-induced mortality. We conclude that strong trade-offs between xylem allocation to fiber walls, fiber lumen, and axial parenchyma drive drought resistance in neotropical trees. Moreover, the benefits of xylem allocation to axial parenchyma in wet tropical trees might not outweigh the consequential low embolism resistance under more frequent and severe droughts in a changing climate., (© 2019 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.)

A substantial portion of locally respired CO 2 in stems can be assimilated by chloroplast-containing tissues. Woody tissue photosynthesis (P wt ) therefore plays a major role in the stem carbon balance. To study the impact of P wt on stem carbon cycling along a gradient of water availability, stem CO 2 efflux (E A ), xylem CO 2 concentration ([CO 2 ]), and xylem water potential (Ψ xylem ) were measured in 4-year-old Populus tremula L. trees exposed to drought stress and different regimes of light exclusion of woody tissues. Under well-watered conditions, local P wt decreased E A up to 30%. Axial CO 2 diffusion (D ax ) induced by distant P wt caused an additional decrease in E A of up to 25% and limited xylem [CO 2 ] build-up. Under drought stress, absolute decreases in E A driven by P wt remained stable, denoting that P wt was not affected by drought. At the end of the dry period, when transpiration was low, local P wt and D ax offset 20% and 10% of stem respiration on a daily basis, respectively. These results highlight (a) the importance of P wt for an adequate interpretation of E A measurements and (b) homeostatic P wt along a drought stress gradient, which might play a crucial role to fuel stem metabolism when leaf carbon uptake and phloem transport are limited., (© 2019 John Wiley & Sons Ltd.)

Wood density (WD) relates to important tree functions such as stem mechanics and resistance against pathogens. This functional trait can exhibit high intraindividual variability both radially and vertically. With the rise of LiDAR-based methodologies allowing nondestructive tree volume estimations, failing to account for WD variations related to tree function and biomass investment strategies may lead to large systematic bias in AGB estimations. Here, we use a unique destructive dataset from 822 trees belonging to 51 phylogenetically dispersed tree species harvested across forest types in Central Africa to determine vertical gradients in WD from the stump to the branch tips, how these gradients relate to regeneration guilds and their implications for AGB estimations. We find that decreasing WD from the tree base to the branch tips is characteristic of shade-tolerant species, while light-demanding and pioneer species exhibit stationary or increasing vertical trends. Across all species, the WD range is narrower in tree crowns than at the tree base, reflecting more similar physiological and mechanical constraints in the canopy. Vertical gradients in WD induce significant bias (10%) in AGB estimates when using database-derived species-average WD data. However, the correlation between the vertical gradients and basal WD allows the derivation of general correction models. With the ongoing development of remote sensing products providing 3D information for entire trees and forest stands, our findings indicate promising ways to improve greenhouse gas accounting in tropical countries and advance our understanding of adaptive strategies allowing trees to grow and survive in dense rainforests.

Under the effect of disturbances, like unbalanced stem, but also during normal development, poplar trees can develop a specific secondary xylem, called "tension wood" (TW), which is easily identifiable by the presence of a gelatinous layer in the secondary cell walls (SCW) of the xylem fibers. Since TW formation was mainly performed on 2-year-old poplar models, an in vitro poplar that produces gelatinous fibers (G-fibers) while offering the same experimental advantages as herbaceous plants has been developed. Using specific cell wall staining techniques, wood structural features and lignin/cellulose distribution were both detailed in cross-sections obtained from the curved stem part of in vitro poplars. A supposed delay in the SCW lignification process in the G-fibers, along with the presence of a G-layer, could be observed in the juvenile plants. Moreover, in this G-layer, the immunolabeling of various polymers carried out in the SCW of TW has allowed detecting crystalline cellulose, arabinogalactans proteins, and rhamnogalacturonans I; however, homogalacturonans, xylans, and xyloglucans could not be found. Interestingly, extensins were detected in this typical adaptative or stress-induced structure. These observations were corroborated by a quantitation of the immunorecognized polymer distribution using gold particle labeling. In conclusion, the in vitro poplar model seems highly convenient for TW studies focusing on the implementation of wall polymers that provide the cell wall with greater plasticity in adapting to the environment.

Biomass and area ratios between leaves, stems and roots regulate many physiological and ecological processes. The Huber value H v (sapwood area/leaf area ratio) is central to plant water balance and drought responses. However, its coordination with key plant functional traits is poorly understood, and prevents developing trait-based prediction models. Based on theoretical arguments, we hypothesise that global patterns in H v of terminal woody branches can be predicted from variables related to plant trait spectra, that is plant hydraulics and size and leaf economics. Using a global compilation of 1135 species-averaged H v , we show that H v varies over three orders of magnitude. Higher H v are seen in short small-leaved low-specific leaf area (SLA) shrubs with low K s in arid relative to tall large-leaved high-SLA trees with high K s in moist environments. All traits depend on climate but climatic correlations are stronger for explanatory traits than H v . Negative isometry is found between H v and K s , suggesting a compensation to maintain hydraulic supply to leaves across species. This work identifies the major global drivers of branch sapwood/leaf area ratios. Our approach based on widely available traits facilitates the development of accurate models of above-ground biomass allocation and helps predict vegetation responses to drought., (© 2019 The Authors. New Phytologist © 2019 New Phytologist Trust.)

The vast majority of measurements in the field of plant hydraulics have been on small-diameter branches from woody species. These measurements have provided considerable insight into plant functioning, but our understanding of plant physiology and ecology would benefit from a broader view, because branch hydraulic properties are influenced by many factors. Here, we discuss the influence that other components of the hydraulic network have on branch vulnerability to embolism propagation. We also modelled the impact of changes in the ratio of root-to-leaf areas and soil texture on vulnerability to hydraulic failure along the soil-to-leaf continuum and showed that hydraulic function is better maintained through changes in root vulnerability and root-to-leaf area ratio than in branch vulnerability. Differences among species in the stringency with which they regulate leaf water potential and in reliance on stored water to buffer changes in water potential also affect the need to construct embolism resistant branches. Many approaches, such as measurements on fine roots, small individuals, combining sap flow and psychrometry techniques, and modelling efforts, could vastly improve our understanding of whole-plant hydraulic functioning. A better understanding of how traits are coordinated across the whole plant will improve predictions for plant function under future climate conditions., (© 2019 John Wiley & Sons Ltd.)

The evolution of secondary (insular) woodiness and the rapid disparification of plant growth forms associated with island radiations show intriguing parallels between oceanic islands and tropical alpine sky islands. However, the evolutionary significance of these phenomena remains poorly understood and the focus of debate. We explore the evolutionary dynamics of species diversification and trait disparification across evolutionary radiations in contrasting island systems compared with their nonisland relatives. We estimate rates of species diversification, growth form evolution and phenotypic space saturation for the classical oceanic island plant radiations - the Hawaiian silverswords and Macaronesian Echium - and the well-studied sky island radiations of Lupinus and Hypericum in the Andes. We show that secondary woodiness is associated with dispersal to islands and with accelerated rates of species diversification, accelerated disparification of plant growth forms and occupancy of greater phenotypic trait space for island clades than their nonisland relatives, on both oceanic and sky islands. We conclude that secondary woodiness is a prerequisite that could act as a key innovation, manifest as the potential to occupy greater trait space, for plant radiations on island systems in general, further emphasizing the importance of combinations of clade-specific traits and ecological opportunities in driving adaptive radiations., (© 2019 The Authors. New Phytologist © 2019 New Phytologist Trust.)

Living vegetation volume (LVV), one of the most difficult tree parameters to calculate, is among the most important factors that indicates the biological characteristics and ecological functions of the crown. Obtaining precise LVV estimates is, however, challenging task because the irregularities of many crown shapes are difficult to capture using standard forestry field equipment. Terrestrial light detection and ranging (T-LiDAR) can be used to record the three-dimensional structures of trees. The primary branches of Larix olgensis and Quercus mongolica in the Qingyuan Experimental Station of Forest Ecology at the Chinese Academy of Sciences (CAS) were taken as the research objects. A new rapid LVV estimation method called the filling method was proposed in this paper based on a T-LiDAR point cloud. In the proposed method, the branch point clouds are divided into leaf points and wood points. We used RiSCAN PRO 1.64 to manually separate the leaf points and wood points under careful visual inspection, and calculated that leaf points and wood points accounted for 91% and 9% of the number of the point clouds of branches. Then, the equation LVV = V1N (where N is the number of leaf points, and V1 is cube size) is used to calculate LVV. When the laser transmission frequency is 300,000 points/second and the point cloud is diluted to 30% using the octree method, the point cloud can be replaced by a cube (V1) of 6.11 cm3 to fill the branch space. The results showed that good performance for this approach, the measuring accuracy for L. olgensis and Q. mongolica at the levels of α = 0.05 and α = 0.01, respectively (94.35%, 90.01% and 91.99%, 85.63%, respectively). The results suggest that the proposed method can be conveniently used to calculate the LVV of coniferous and broad-leaf species under specific scanning settings. This work is explorative because hypotheses or a theoretical framework have not been previously defined. Rather, we would like to contribute to the formation of hypotheses as a background for further studies., Competing Interests: The authors have declared that no competing interests exist.

An investigation of simultaneous dynamic mass and length change measurement for wood is presented. In contrast to the equilibrium in moisture content and swelling and shrinking, where extensive data exists for different wood species, less information is available for the dynamics of moisture changes in direct comparison to the related dimensional changes during the sorption process. This is due to a lack of methods. A gravimetric sorption system, equipped with a high resolution camera and an automated image evaluation, is used to examine simultaneous effects of water vapour sorption dynamics and dimensional change. This method proves a strong correlation between mass and dimensional change, which is in contrast to other investigations. Equilibrium moisture content as well as swelling and shrinking data is in good agreement with literature and manual measurements. The method enables the possibility to determine swelling and shrinking values in-situ without disturbing the targeted climatic conditions. The system is applicable for the investigation of natural wood, modified wood, wood composites or other lignocellulosic materials.

Reactivation of axial water flow in ring-porous species is a complex process related to stem water content and developmental stage of both earlywood-vessel and leaf formation. Yet empirical evidence with non-destructive methods on the dynamics of water flow resumption in relation to these mechanisms is lacking. Here we combined in vivo magnetic resonance imaging and wood-anatomical observations to monitor the dynamic changes in stem water content and flow during spring reactivation in 4-year-old pedunculate oaks (Quercus robur L.) saplings. We found that previous year latewood vessels and current year developing earlywood vessels form a functional unit for water flow during growth resumption. During spring reactivation, water flow shifted from latewood towards the new earlywood, paralleling the formation of earlywood vessels and leaves. At leaves' full expansion, volumetric water content of previous rings drastically decreased due to the near-absence of water in fibre tissue. We conclude (i) that in ring-porous oak, latewood vessels play an important hydraulic role for bridging the transition between old and new water-conducting vessels and (ii) that fibre and parenchyma provides a place for water storage., (© The Author(s) 2019. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Xylem conduit diameter (D max ) of woody angiosperm adults scales with plant size and widens from the stem apex downwards. We hypothesized that, notwithstanding relative growth rate (RGR), growth form or leaf habit, woody seedling conduit D max scales linearly with plant size across species; this scaling should be applicable to all vegetative organs, with consistent conduit widening from leaf via stem to main root and coupling with whole-leaf area and whole-stem xylem area. To test these hypotheses, organ-specific xylem anatomy traits and size-related traits in laboratory-grown seedlings were analyzed across 55 woody European species from cool-temperate and Mediterranean climates. As hypothesized, conduit D max of each organ showed similar scaling with plant size and consistent basipetal widening from the leaf midvein via the stem to the main root across species, independently of growth form, RGR and leaf habit. We also found a strong correlation between D max and average leaf area, and between stem xylem area and whole-plant leaf area. We conclude that seedlings of ecologically wide-ranging woody species converge in their allometric scaling of conduit diameters within and across plant organs. These relationships will contribute to modeling of water transport in woody vegetation that accounts for the whole life history from the trees' regeneration phase to adulthood., (© 2019 The Authors. New Phytologist © 2019 New Phytologist Trust.)

Hydraulic safety and efficiency have become the central concept of the interpretation of the structure and function of vessels and their interconnections. Plants form an appropriate xylem network structure to maintain a balance of hydraulic safety vs efficiency. The term 'tracheid bridge' is used to describe a possible pathway of water transport between neighboring vessels via tracheids, and this pathway could also provide increased safety against embolisms. However, the only physiological study of such a structure thus far has been in Hippophae rhamnoides Linn. To test the function of tracheid bridges, this research examined four species that have relatively long and solitary vessels, which are two of the criteria for efficient tracheid bridges. Tracheids contributed less than 2.2% of the total conductance of the vessels in these species, but in theory, tracheids could serve as very efficient transport connector pathways that may or may not make direct vessel-to-vessel contact via pit fields between adjacent vessels. In some species, tracheid bridges may represent the dominant pathway for water flow between vessels, whereas in other species, tracheid bridges may be sub-dominant or virtually nil. Broader searches of woody taxa are needed to reveal the functional importance of tracheid bridges in vascular plants., (© The Author(s) 2019. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

The concept of growth rings is little discussed in the literature and their treatment remains somewhat confusing in terms of the diversity of structures described. This situation has a major impact on the study of growth rings in tropical species, in which variations and complexity are greater and accuracy of identification less good. The rigid conceptual delimitations used by dendrochronologists and wood anatomists of temperate regions cannot be applied to the study of growth rings in most tropical species, which has led to neglect of this subject. With the objective of discussing the concept of growth rings, the present study consists of a survey of anatomical, periodicity, causal and evolutionary aspects of the treatment of these structures as evidenced in previous studies, and the evaluation of their application and limitations to the development of this concept. Anatomical aspects arise through radial integrity, or the presence of early and late wood; the degree of distinction of the rings, which may vary from well to poorly defined; and tangential continuity, meaning that rings may form a complete circle in the transverse section of the trunk, or instead be tangentially discontinuous, lens-shaped or in wedges. In addition there are a diversity of anatomical markers which enable us to recognize growth rings. Regarding periodicity, the rings may be annual, infra-annual or supra-annual. Causal aspects include genetic, endogenous and environmental components. Evolutionary aspects continue to be insufficiently investigated, and although most comparative studies treat growth rings recognized by different markers as though they were a single character, it remains questionable whether they can be regarded as homologous or the result of homoplasy. We conclude that the elaboration of a robust but broad definition which can include all the variation observed in growth rings of tropical species is a complex task, which is only possible by overcoming the restrictions adopted by dendrochronologists and wood anatomists of temperate regions for whom growth rings are essentially annual, strongly demarcated, tangentially continuous and restricted to the most pronounced markers., (© 2019 Cambridge Philosophical Society.)