Ecological stress memory in wood architecture of two Neotropical hickory species from central-eastern Mexico.

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.

Seasonal variation of vessel pits in sapwood: microscopical analyses of the morphology and chemical components of pit membrane encrustations in Fraxinus mandschurica.

BACKGROUND AND AIMS: Pit pairs and their filter-like partition, i.e. pit membranes, play important roles as water pathways, barriers and regulators in the water-conducting system of angiosperms. In Fraxinus species, the intervessel and vessel-parenchyma pit membranes in sapwood are normally encrusted during winter. Although these encrustations inevitably influence the performance of pits, their properties and functions remain unclear. This study aimed to reveal the morphological and chemical characteristics of encrustations in F. mandshurica in order to deepen understanding of the seasonal encrustation of pit membranes. METHODS: Seasonal and positional variations in the presence and morphology of encrustations were examined by field-emission scanning electron microscopy (FE-SEM). Cryo-FE-SEM for freeze-fixed greenwood samples was conducted to clarify whether encrustations were present in living trees. Chemical components were examined by histochemical staining using light and electron microscopy, immunofluorescence labelling and ultraviolet microspectroscopy. KEY RESULTS: Encrustations began to deposit in autumn before leaf senescence and disappeared in spring before bud flushing. They infiltrated within the pit membranes, which suggested that they severely limit the permeation of pits. The encrustations differed in morphology among positions: they entirely filled the pit chambers in latewood, while they covered the pit membranes in earlywood. The encrustations were similarly observed in the samples that were freeze-fixed immediately after collection, indicating that they are present in living trees. The encrustations contained polysaccharides, including xyloglucan and homogalacturonan, and phenolic compounds, possibly including flavonoids and coumarins. These chemical components were also detected in droplets found in the latewood vessels with the encrustations, suggesting that the materials constituting encrustations were supplied through the vessel lumens. CONCLUSIONS: Encrustations undoubtedly cover the pit membranes in living F. mandshurica trees in winter and their morphology and chemical composition indicate that they are impermeable, have positional differences in function and are characterized by elaborate deposition/removal processes.

The potential of X-ray computed tomography for xylological and dendrochronological analyses of Egyptian mummy labels.

X-ray computed tomography (XRCT) imaging allows non-destructive visualization of the structure of various materials. Applied to wooden objects, it allows determination of their morphologies or manufacturing techniques, but also measurement of growth ring widths. We have applied XRCT to a selection of 38 mummy labels. This funerary furniture, made up of endemic or imported tree species, has survived thanks to environmental conditions in very large quantities in regions in Middle and Upper Egypt and is featured now in museum collections across the globe. Mummy labels thus represent a unique and abundant data source to build floating or absolutely dated dendrochronological chronologies for this period. Here we discuss the possible contributions and limitations of XRCT for the analysis of these artifacts and show that the approach allows identification of discriminating markers for the identification of certain species on the transverse plane, but that the insufficient resolution of the tangential and radial planes normally prevents formal identification of species. By contrast, XRCT undeniably enhances the visibility of toolmarks (in terms of numbers and depth), and thereby allows highlighting marks that remain invisible to the naked eye; XRCT also provides key insights into cutting methods and the calibers used and yields new information on silvicultural practices and the knowhow of Egyptian craftsmen. Finally, the measurement of ring widths on XRCT imagery is also more accurate than what can be achieved by traditional dendrochronological measurements, especially in the case of cuts realized on a slab. The approach also confirms the limited potential of local broadleaved species for dendrochronological approaches due to unreadable or poorly visible tree rings and mostly short tree-ring sequences.

Advancing fine branch biomass estimation with lidar and structural models.

BACKGROUND AND AIMS: Lidar is a promising tool for fast and accurate measurements of trees. There are several approaches to estimate above-ground woody biomass using lidar point clouds. One of the most widely used methods involves fitting geometric primitives (e.g. cylinders) to the point cloud, thereby reconstructing both the geometry and topology of the tree. However, current algorithms are not suited for accurate estimation of the volume of finer branches, because of the unreliable point dispersions from, for example, beam footprint compared to the structure diameter. METHOD: We propose a new method that couples point cloud-based skeletonization and multi-linear statistical modelling based on structural data to make a model (structural model) that accurately estimates the above-ground woody biomass of trees from high-quality lidar point clouds, including finer branches. The structural model was tested at segment, axis and branch level, and compared to a cylinder fitting algorithm and to the pipe model theory. KEY RESULTS: The model accurately predicted the biomass with 1.6 % normalized root mean square error (nRMSE) at the segment scale from a k-fold cross-validation. It also gave satisfactory results when scaled up to the branch level with a significantly lower error (13 % nRMSE) and bias (-5 %) compared to conventional cylinder fitting to the point cloud (nRMSE: 92 %, bias: 82 %), or using the pipe model theory (nRMSE: 31 %, bias: -27 %). The model was then applied to the whole-tree scale and showed that the sampled trees had more than 1.7 km of structures on average and that 96 % of that length was coming from the twigs (i.e. <5 cm diameter). Our results showed that neglecting twigs can lead to a significant underestimation of tree above-ground woody biomass (-21 %). CONCLUSIONS: The structural model approach is an effective method that allows a more accurate estimation of the volumes of smaller branches from lidar point clouds. This method is versatile but requires manual measurements on branches for calibration. Nevertheless, once the model is calibrated, it can provide unbiased and large-scale estimations of tree structure volumes, making it an excellent choice for accurate 3D reconstruction of trees and estimating standing biomass.

Automating Wood Species Detection and Classification in Microscopic Images of Fibrous Materials with Deep Learning.

We have developed a methodology for the systematic generation of a large image dataset of macerated wood references, which we used to generate image data for nine hardwood genera. This is the basis for a substantial approach to automate, for the first time, the identification of hardwood species in microscopic images of fibrous materials by deep learning. Our methodology includes a flexible pipeline for easy annotation of vessel elements. We compare the performance of different neural network architectures and hyperparameters. Our proposed method performs similarly well to human experts. In the future, this will improve controls on global wood fiber product flows to protect forests.

Climate of origin shapes variations in wood anatomical properties of 17 Picea species.

BACKGROUND: Variations in hydraulic conductivity may arise from species-specific differences in the anatomical structure and function of the xylem, reflecting a spectrum of plant strategies along a slow-fast resource economy continuum. Spruce (Picea spp.), a widely distributed and highly adaptable tree species, is crucial in preventing soil erosion and enabling climate regulation. However, a comprehensive understanding of the variability in anatomical traits of stems and their underlying drivers in the Picea genus is currently lacking especially in a common garden. RESULTS: We assessed 19 stem economic properties and hydraulic characteristics of 17 Picea species grown in a common garden in Tianshui, Gansu Province, China. Significant interspecific differences in growth and anatomical characteristics were observed among the species. Specifically, xylem hydraulic conductivity (Ks) and hydraulic diameter exhibited a significant negative correlation with the thickness to span ratio (TSR), cell wall ratio, and tracheid density and a significant positive correlation with fiber length, and size of the radial tracheid. PCA revealed that the first two axes accounted for 64.40% of the variance, with PC1 reflecting the trade-off between hydraulic efficiency and mechanical support and PC2 representing the trade-off between high embolism resistance and strong pit flexibility. Regression analysis and structural equation modelling further confirmed that tracheid size positively influenced Ks, whereas the traits DWT, D_r, and TSR have influenced Ks indirectly. All traits failed to show significant phylogenetic associations. Pearson's correlation analysis demonstrated strong correlations between most traits and longitude, with the notable influence of the mean temperature during the driest quarter, annual precipitation, precipitation during the wettest quarter, and aridity index. CONCLUSIONS: Our results showed that xylem anatomical traits demonstrated considerable variability across phylogenies, consistent with the pattern of parallel sympatric radiation evolution and global diversity in spruce. By integrating the anatomical structure of the stem xylem as well as environmental factors of origin and evolutionary relationships, our findings provide novel insights into the ecological adaptations of the Picea genus.

Multimodal imaging analysis in silver fir reveals coordination in cellulose and lignin deposition.

Despite lignin being a key component of wood, the dynamics of tracheid lignification are generally overlooked in xylogenesis studies, which hampers our understanding of environmental drivers and blurs the interpretation of isotopic and anatomical signals stored in tree rings. Here, we analyzed cell wall formation in silver fir (Abies alba Mill.) tracheids to determine if cell wall lignification lags behind secondary wall deposition. For this purpose, we applied a multimodal imaging approach combining transmitted light microscopy (TLM), confocal laser scanning microscopy (CLSM), and confocal Raman microspectroscopy (RMS) on anatomical sections of wood microcores collected in northeast France on 11 dates during the 2010 growing season. Wood autofluorescence after laser excitation at 405 and 488 nm associated with the RMS scattering of lignin and cellulose, respectively, which allowed identification of lignifying cells (cells showing lignified and nonlignified wall fractions at the same time) in CLSM images. The number of lignifying cells in CLSM images mirrored the number of wall-thickening birefringent cells in polarized TLM images, revealing highly synchronized kinetics for wall thickening and lignification (similar timings and durations at the cell level). CLSM images and RMS chemical maps revealed a substantial incorporation of lignin into the wall at early stages of secondary wall deposition. Our results show that most of the cellulose and lignin contained in the cell wall undergo concurrent periods of deposition. This suggests a strong synchronization between cellulose and lignin-related features in conifer tree-ring records, as they originated over highly overlapped time frames.

Stretched sapwood, ultra-widening permeability and ditching da Vinci: revising models of plant form and function.

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.

Variations in wood anatomy in Afrotropical trees with a particular emphasis on radial and axial parenchyma.

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.

Analyses of high spatial resolution datasets identify genes associated with multi-layered secondary cell wall thickening in Pinus bungeana.

BACKGROUND AND AIMS: Secondary cell wall (SCW) thickening is a major cellular developmental stage determining wood structure and properties. Although the molecular regulation of cell wall deposition during tracheary element differentiation has been well established in primary growth systems, less is known about the gene regulatory processes involved in the multi-layered SCW thickening of mature trees. METHODS: Using third-generation [long-read single-molecule real-time (SMRT)] and second-generation [short-read sequencing by synthesis (SBS)] sequencing methods, we established a Pinus bungeana transcriptome resource with comprehensive functional and structural annotation for the first time. Using these approaches, we generated high spatial resolution datasets for the vascular cambium, xylem expansion regions, early SCW thickening, late SCW thickening and mature xylem tissues of 71-year-old Pinus bungeana trees. KEY RESULTS: A total of 79 390 non-redundant transcripts, 31 808 long non-coding RNAs and 5147 transcription factors were annotated and quantified in different xylem tissues at all growth and differentiation stages. Furthermore, using this high spatial resolution dataset, we established a comprehensive transcriptomic profile and found that members of the NAC, WRKY, SUS, CESA and LAC gene families are major players in early SCW formation in tracheids, whereas members of the MYB and LBD transcription factor families are highly expressed during late SCW thickening. CONCLUSIONS: Our results provide new molecular insights into the regulation of multi-layered SCW thickening in conifers. The high spatial resolution datasets provided can serve as important gene resources for improving softwoods.

An open-source machine-learning approach for obtaining high-quality quantitative wood anatomy data from E. grandis and P. radiata xylem.

Quantitative wood anatomy is a subfield in dendrochronology that requires effective open-source image analysis tools. In this research, the bioimage analysis software QuPath (v0.4.4) is introduced as a candidate for accurately quantifying the cellular properties of the xylem in an automated manner. Additionally, the potential of QuPath to detect the transition of early- to latewood tracheids over the growing season was evaluated to assess a potential application in dendroecological studies. Various algorithms in QuPath were optimized to quantify different xylem cell types in Eucalyptus grandis and the transition of early- to latewood tracheids in Pinus radiata. These algorithms were coded into cell detection scripts for automatic quantification of stem microsections and compared to a manually curated method to assess the accuracy of the cell detections. The automatic cell detection approach, using QuPath, has been validated to be reproducible with an acceptable error when assessing fibers, vessels, early- and latewood tracheids. However, further optimization for parenchyma is still required. This proposed method developed in QuPath provides a scalable and accurate approach for quantifying anatomical features in stem microsections. With minor amendments to the detection and classification algorithms, this strategy is likely to be viable in other plant species.

Impact of intra-annual wood density fluctuation on tree hydraulic function: insights from a continuous monitoring approach.

Climate change significantly impacts global forests, leading to tree decline and dieback. To cope with climate change, trees develop several functional traits, such as intra-annual density fluctuations (IADFs) in tree rings. The formation of these traits facilitates trees to optimize resource allocation, allowing them to withstand periods of stress and eventually recover when the conditions become favourable again. This study focuses on a Pinus pinaster Aiton forest in a warm, drought-prone Mediterranean area, comparing two growing seasons with different weather patterns. The innovative continuous monitoring approach used in this study combines high-resolution monitoring of sap flow (SF), analysis of xylogenesis and quantitative wood anatomy. Our results revealed the high plasticity of P. pinaster in water use and wood formation, shedding light on the link between IADFs and tree conductance. Indeed, the capacity to form large cells in autumn (as IADFs) improves the total xylem hydraulic conductivity of this species. For the first time, a continuous SF measurement system captured the dynamics of bimodal SF during the 2022 growing season in conjunction with the bimodal growth pattern observed through xylogenesis monitoring. These results highlight the intricate interplay between environmental conditions, water use, wood formation and tree physiology, providing valuable insights into the acclimation mechanisms employed by P. pinaster to cope with weather fluctuations.

Formation of xylem tissues and secondary cell walls is diminished by severe and consecutive insect defoliation.

PREMISE: Insect defoliation of trees causes unusual changes to wood anatomy and slows radial growth that decreases tree value; however, the characteristics of these anatomical changes in hardwoods remain unclear. The aim of this study was to characterize the anatomy and histochemistry of the wood in trunks of Betula maximowicziana trees after severe insect defoliation. METHODS: Secondary xylem tissues were sampled from trunks that had been defoliated by Caligula japonica at Naie and Furano in central Hokkaido during 2006-2012, then cross-dated and examined microscopically and stained histochemically to characterize anatomical and chemical changes in the cells. RESULTS: White rings with thin-walled wood fibers and greatly reduced annual ring width in the subsequent year were observed in samples from both sites. From these results, the year that the white rings formed was determined, and severe defoliation was confirmed to trigger white ring formation. The characteristics may prove useful to detect the formation year of white rings. Scanning electron microscopy and histochemical analyses of the white rings indicated that the thickness of the S2 layer in the wall of wood fiber cells decreased, but xylan and lignin were still deposited in the cell walls of wood fibers. However, the walls of the fibers rethickened after the defoliation. CONCLUSIONS: Our results suggest that B. maximowicziana responds to a temporary lack of carbon inputs due to insect defoliation by regulating the thickness of the S2 layer of the cell wall of wood fibers. For B. maximowicziana, insect defoliation late in the growing season has serious deleterious effects on wood formation and radial growth.

Toward a better understanding of angiosperm xylogenesis: a new method for a cellular approach.

The kinetics of wood formation in angiosperms are largely unknown because their complex xylem anatomy precludes using the radial position of vessels and fibers to infer their time of differentiation. We analyzed xylogenesis in ring-porous ash (Fraxinus angustifolia) and diffuse-porous beech (Fagus sylvatica) over 1 yr and proposed a novel procedure to assess the period of vessel and fiber enlargement using a referential radial file (RRF). Our approach captured the dynamics of wood formation and provided a robust estimation of the kinetics of vessel and fiber enlargement. In beech, fibers and vessels had a similar duration of enlargement, decreasing from 14 to 5 d between April and July. In ash, wide vessels formed in April enlarged at a rate of 27 × 103 µm2 d-1 , requiring half the time of contemporary fibers (6 vs 12 d), and less time than the narrower vessels (14 d) formed in May. These findings reveal distinct cell-type-dependent mechanisms for differentiation in diffuse-porous and ring-porous trees, enhancing our understanding of angiosperm wood cell kinetics. Our approach presents an effective method for investigating angiosperm wood formation and provides a more accurate representation of vessel and fiber morphogenesis in wood formation models.

Addressing controversies in the xylem embolism resistance-vessel diameter relationship.

Although xylem embolism is a key process during drought-induced tree mortality, its relationship to wood anatomy remains debated. While the functional link between bordered pits and embolism resistance is known, there is no direct, mechanistic explanation for the traditional assumption that wider vessels are more vulnerable than narrow ones. We used data from 20 temperate broad-leaved tree species to study the inter- and intraspecific relationship of water potential at 50% loss of conductivity (P50 ) with hydraulically weighted vessel diameter (Dh ) and tested its link to pit membrane thickness (TPM ) and specific conductivity (Ks ) on species level. Embolism-resistant species had thick pit membranes and narrow vessels. While Dh was weakly associated with TPM , the P50 -Dh relationship remained highly significant after accounting for TPM . The interspecific pattern between P50 and Dh was mirrored by a link between P50 and Ks , but there was no evidence for an intraspecific relationship. Our results provide robust evidence for an interspecific P50 -Dh relationship across our species. As a potential cause for the inconsistencies in published P50 -Dh relationships, our analysis suggests differences in the range of trait values covered, and the level of data aggregation (species, tree or sample level) studied.

A safe breeding ground: genetically improved maritime pine for growth and stem form has more efficient but not more vulnerable xylem.

Through repeated cycles of selection and recombination, tree breeding programs deliver genetically improved varieties for a range of target characteristics such as biomass production, stem form, resistance to biotic stresses, wood properties, etc. However, in the context of increased drought and heat waves, it is not yet known whether growth performance will impede drought resistance. To address this question, we compared the hydraulic properties, such as hydraulic efficiency and hydraulic safety, in four varieties over successive varieties of genetically improved maritime pines (i.e., Pinus pinaster Aït.) for growth and stem form. We measured 22 functional traits related to hydraulic efficiency, hydraulic safety, xylem anatomy and wood density. We found that improved varieties presented higher hydraulic conductivity with larger tracheid lumen size and tracheid lumen fraction, and smaller wall thickness reinforcement and tracheid density, but not at the cost of reduced embolism resistance. The reported absence of trade-off between hydraulic conductivity and embolism resistance is a strong asset to improve biomass productivity, through increased hydraulic efficiency, without impacting drought resistance, and should enable new maritime pine varieties to cope with a drier climate. Our study is one of the first to reveal the hydraulic mechanisms over successive varieties of genetic improvement for tree growth. It provides guidelines for sustainable forest management through breeding for other forest tree species.

Tree-ring anatomy of Pinus cembra trees opens new avenues for climate reconstructions in the European Alps.

Tree rings form the backbone of high-resolution palaeoclimatology and represent one of the most frequently used proxy to reconstruct climate variability of the Common Era. In the European Alps, reconstructions were often based on tree-ring width (TRW) and maximum latewood density (MXD) series, with a focus on European larch. By contrast, only a very limited number of dendroclimatic studies exists for long-lived, multi-centennial Pinus cembra, despite the widespread occurrence of the species at treeline sites across the European Alps. This lack of reconstructions can be ascribed to the difficulties encountered in past studies in extracting a robust climate signal from TRW and MXD chronologies. In this study, we tested various wood anatomical parameters from P. cembra as proxies for the reconstruction of past air temperatures. To this end, we measured anatomical cell parameters and TRW of old-growth trees from the God da Tamangur forest stand, known for being the highest pure, and continuous P. cembra forest in Europe. We demonstrate that several wood anatomical parameters allow robust reconstruction of past temperature variability at annual to multidecadal timescales. Best results are obtained with maximum latewood radial cell wall thickness (CWTrad) measured at 40 µm radial band width. Over the 1920-2017 period, the CWTrad chronology explains 62 % and >80 % of interannual and decadal variability of air temperatures during a time window corresponding roughly with the growing season. These values exceed those found in past work on P. cembra and even exceed the values reported for MXD chronologies built with L. decidua and hitherto considered the gold standard for dendroclimatic reconstructions in the European Alps. The wood anatomical analysis of P. cembra records therefore unveils a dormant potential and opens new avenues for a species that has been considered unsuitable for climate reconstructions so far.

The vessel wall thickness-vessel diameter relationship across woody angiosperms.

PREMISE: Comparative anatomy is necessary to identify the extremes of combinations of functionally relevant structural traits, to ensure that physiological data cover xylem anatomical diversity adequately, and thus achieve a global understanding of xylem structure-function relations. A key trait relationship is that between xylem vessel diameter and wall thickness of both the single vessel and the double vessel+adjacent imperforate tracheary element (ITE). METHODS: We compiled a comparative data set with 1093 samples, 858 species, 350 genera, 86 families, and 33 orders. We used broken linear regression and an algorithm to explore changes in parameter values from linear regressions using subsets of the data set to identify a threshold, at 90-µm vessel diameter, in the wall thickness-diameter relationship. RESULTS: Below 90 µm diameter for vessels, virtually any wall thickness could be associated with virtually any diameter. Below this threshold, selection is free to favor a very wide array of combinations, such as very thick walls and narrow vessels in ITE-free herbs, or very thin-walled, wide vessels in evergreen dryland pioneers. Above 90 µm, there was a moderate positive relationship. CONCLUSIONS: Our analysis shows that the space of vessel wall thickness-diameter combinations is very wide, with selection apparently eliminating individuals with vessel walls "too thin" for their diameter. Most importantly, our survey revealed poorly studied plant hydraulic syndromes (functionally significant trait combinations). These data suggest that the full span of trait combinations, and thus the minimal set of hydraulic syndromes requiring study to span woody plant functional diversity adequately, remains to be documented.

The 2018 hot drought pushed conifer wood formation to the limit of its plasticity: Consequences for woody biomass production and tree ring structure.

Hot droughts are expected to increase in Europe and disturb forest ecosystem functioning. Wood formation of trees has the potential to adapt to those events by compensatory mechanisms between the rates and durations of tracheid differentiation to form the typical pattern of vital wood anatomical structures. We monitored xylogenesis and measured wood anatomy of mature silver fir (Abies alba Mill.) and Scots pine (Pinus sylvestris L.) trees along an elevational gradient in the Black Forest during the hot drought year of 2018. We assessed the kinetics of tracheid differentiation and the final tracheid dimensions and quantified the relationship between rates and durations of cell differentiation over the growing season. Cell differentiation kinetics were decoupled, and temperature and water availability signals were imprinted in the tree ring structure. The sudden decline in woody biomass production provided evidence for a disruption in carbon sequestration processes due to heat and drought stress. Growth processes of Scots pine (pioneer species) were mainly affected by the spring drought, whereas silver fir (climax species) growth processes were more disturbed by the summer drought. Our study provides novel insights on the plasticity of wood formation and carbon allocation in temperate conifer tree species in response to extreme climatic events.

Cutting tree rings into time slices: how intra-annual dynamics of wood formation help decipher the space-for-time conversion.

Tree-ring anatomy, microdensity and isotope records provide valuable intra-annual information. However, extracting signals at that scale is challenged by the complexity of xylogenesis, where two major processes - cell enlargement and wall thickening - occur at different times and rates. We characterized the space-for-time association in the tree rings of three conifer species by examining the duration, overlapping, inter-tree synchronicity and interannual stability during cell enlargement and wall thickening across regular tree-ring sectors (portions of equal tangential width). The number of cells and cell differentiation rates determined the duration of sector formation, which augmented more rapidly throughout the ring for wall thickening than for enlargement. Increasing the number of sectors above c. 15 had a limited effect on improving time resolution because consecutive sector formation overlapped greatly in time, especially in narrow rings and during wall thickening. Increasing the number of sectors also resulted in lower synchronicity and stability of intermediate-sector enlargement, whereas all sectors showed high synchronicity and stability during wall thickening. Increasing the number of sectors had a stronger effect on enhancing time-series resolution for enlargement- than for wall-thickening-related traits, which would nevertheless produce more reliable intra-annual chronologies as a result of the more similar calendars across trees and years in wall thickening.