Search

Sap exudation is the process whereby trees such as sugar (Acer saccharum Marsh.) and red maple (Acer rubrum L.) generate unusually high positive stem pressure in response to repeated cycles of freeze and thaw. This elevated xylem pressure permits the sap to be harvested over a period of several weeks and hence is a major factor in the viability of the maple syrup industry. The extensive literature on sap exudation documents competing hypotheses regarding the physical and biological mechanisms that drive positive pressure generation in maple, but to date, relatively little effort has been expended on devising mathematical models for the exudation process. In this paper, we utilize an existing model of Graf et al. (J Roy Soc Interface 12:20150665, 2015) that describes heat and mass transport within the multiphase gas–liquid–ice mixture in the porous xylem tissue. The model captures the inherent multiscale nature of xylem transport by including phase change and osmotic transport in wood cells on the microscale, which is coupled to heat transport through the tree stem on the macroscale. A parametric study based on simulations with synthetic temperature data identifies the model parameters that have greatest impact on stem pressure build-up. Measured daily temperature fluctuations are then used as model inputs and the resulting simulated pressures are compared directly with experimental measurements taken from mature red and sugar maple stems during the sap harvest season. The results demonstrate that our multiscale freeze–thaw model reproduces realistic exudation behavior, thereby providing novel insights into the specific physical mechanisms that dominate positive pressure generation in maple trees. [ABSTRACT FROM AUTHOR]

The critical role of auxin on secondary vascular development in woody plants has been demonstrated. The concentration gradient of endogenous indole-3-acetic acid and the cellular and molecular pathways contributing to the auxin-directed vascular organization and wood growth have been uncovered in recent decades. However, our understanding of the roles and regulations of auxin influx in wood formation in trees remains limited. Here, we reported that a microRNA, miR7833, participates in the negative regulation of stem cambial cell division and secondary xylem development in Populus tomentosa. The miR7833 is mainly expressed in the vascular cambium during stem radical growth and specifically targets and represses two AUX/LAX family auxin influx carriers, AUX5 and AUX6 , in poplar. We further revealed that poplar AUX6 , the most abundant miR7833 target in the stem, is preferentially enriched in the developing xylem and is a positive regulator for cell division and differentiation events during wood formation. Moreover, inhibition of auxin influx carriers by 1-naphthoxyacetic acids abolished the regulatory effects of miR7833 and AUX6 on secondary xylem formation in poplar. Our results revealed the essential roles of the miR7833- AUX6 module in regulating cellular events in secondary xylem development and demonstrated an auxin influx-dependent mechanism for wood formation in poplar. [ABSTRACT FROM AUTHOR]

Wood formation is a complex developmental process under the control of multiple levels of regulatory transcriptional network and hormone signals in trees. It is well known that cytokinin (CK) signaling plays an important role in maintaining the activity of the vascular cambium. The CK response factors (CRFs) encoding a subgroup of AP2 transcription factors have been identified to mediate the CK-dependent regulation in different plant developmental processes. However, the functions of CRFs in wood development remain unclear. Here, we characterized the function of PtCRF1, a CRF transcription factor isolated from poplar, in the process of wood formation. The PtCRF1 is preferentially expressed in secondary vasculature, especially in vascular cambium and secondary phloem, and encodes a transcriptional activator. Overexpression of PtCRF1 in transgenic poplar plants led to a significant reduction in the cell layer number of vascular cambium. The development of wood tissue was largely promoted in the PtCRF1 -overexpressing lines, while it was significantly compromised in the CRISPR/Cas9-generated double mutant plants of PtCRF1 and its closest homolog PtCRF2. The RNA sequencing (RNA-seq) and quantitative reverse transcription PCR (RT-qPCR) analyses showed that PtCRF1 repressed the expression of the typical CK-responsive genes. Furthermore, bimolecular fluorescence complementation assays revealed that PtCRF1 competitively inhibits the direct interactions between histidine phosphotransfer proteins and type-B response regulator by binding to PtHP protein. Collectively, these results indicate that PtCRF1 negatively regulates CK signaling and is required for woody cell differentiation in poplar. [ABSTRACT FROM AUTHOR]

A comparative assessment of bud and wood phenology could aid a better understanding of tree growth dynamics. However, the reason for asynchronism or synchronism in leaf and cambial phenology remains unclear. To test the assumption that the temporal relationship between the budburst date and the onset date of wood formation is due to their common or different responses to environmental factors, we constructed a wood phenology dataset from previous literature, and compared it with an existing bud phenology dataset in Europe. We selected three common conifers (Larix decidua Mill. Picea abies (L.) H. Karst. and Pinus sylvestris L.) in both datasets and analyzed 909 records of the onset of wood formation at 47 sites and 238,720 records of budburst date at 3051 sites. We quantified chilling accumulation (CA) and forcing requirement (FR) of budburst and onset of wood formation based on common measures of CA and FR. We then constructed negative exponential CA–FR curves for bud and wood phenology separately. The results showed that the median, variance and probability distribution of CA–FR curves varied significantly between bud and wood phenology for three conifers. The different FR under the same chilling condition caused asynchronous bud and wood phenology. Furthermore, the CA–FR curves manifested that wood phenology was more sensitive to chilling than bud phenology. Thus, the FR of the onset of wood formation increases more than that of budburst under the same warming scenarios, explaining the stronger earlier trends in the budburst date than the onset date of woody formation simulated by the process-based model. Our work not only provides a possible explanation for asynchronous bud and wood phenology from the perspective of organ-specific responses to chilling and forcing, but also develops a phenological model for predicting both bud and wood phenology with acceptable uncertainties. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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 (P 50) ranged from −2.14 to −2.58 MPa. The P 50 and P 88 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 P 50 and P 88 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. [ABSTRACT FROM AUTHOR]

The connections between the primary and secondary growth of trees allows better understanding of the dynamics of carbon sequestration in forest ecosystems. The relationship between primary and secondary growth of trees could change due to the diverging responses of meristems to climate warming. In this study, the bud phenology and radial growth dynamics of Qinghai spruce (Picea crassifolia) in arid and semi-arid areas of China in 2019 and 2020 were weekly monitored to analyze their response to different weather conditions and their links with carbon sink. Xylem anatomical traits (i.e. lumen radial diameter and cell wall thickness) were quantified along cell radial files after the end of xylem lignification to calculate the early-to-latewood transition date. Winter and early spring (January–March) were warmer in 2020 with a colder April compared with 2019. Precipitation in April–June was lower in 2020 than in 2019. In 2019, bud phenology occurred earlier, while the onset of xylem formation and the early-to-latewood transition date were delayed. The duration from the beginning of split bud and exposed shoot to the early-to-latewood transition date was positively correlated with the radial width of earlywood (accounting for ~80% of xylem width) and total xylem width. The longer duration of xylem cell division did not increase xylem cell production and radial width. Moreover, the duration from bud burst to the early-to-latewood transition date in 2020 was negatively linked with early phloem cell production as compared with 2019. Our findings suggest that warm conditions in winter and early spring promote the xylogenesis of Qinghai spruce, but might delay bud burst. However, the xylem width increments largely depend on the duration from bud burst to the start of latewood cell division rather than on the earlier xylogenesis and longer duration of xylem cell differentiation induced by warm conditions. [ABSTRACT FROM AUTHOR]

Bamboo is a highly renewable biomass resource with outstanding ecological, economic and social benefits. However, its lengthy vegetative growth stage and uncertain flowering period have hindered the application of traditional breeding methods. In recent years, significant progress has been made in bamboo breeding. While technical advances in bamboo breeding have been impressive, it is essential to also consider the broader implications we can learn from bamboo's extraordinary features for sustainable development. This review provides an overview of the current status of bamboo breeding technology, including a detailed history of bamboo breeding divided into four eras, a comprehensive map of bamboo germplasm gardens worldwide, with a focus on China, and a summary of available transgenic technologies for gene function verification and genetic improvement. As the demand for bamboo as a sustainable and renewable resource increases continuously, breeding objectives should be focused on enhancing yield, wood properties and adaptability to diverse environmental conditions. In particular, priority should be given to improving fiber length, internode length and wall thickness, as well as regulating lignin and cellulose content for papermaking, substitute for plastic and other applications. Furthermore, we highlight the challenges and opportunities for future research and development in bamboo breeding, including the application of omics technologies, artificial intelligence and the development of new breeding methods. Finally, by integrating the technical advances in bamboo breeding with a discussion of its broader implications for sustainable development, this review provides a comprehensive framework for the development of bamboo industry. [ABSTRACT FROM AUTHOR]

Leaves are the most important photosynthetic organs in most woody plants, but chloroplasts are also found in organs optimized for other functions. However, the actual photosynthetic efficiency of these chloroplasts is still unclear. We analyzed bark and wood chloroplasts of Fraxinus ornus L. saplings. Optical and spectroscopic methods were applied to stem samples and compared with leaves. A sharp light gradient was detected along the stem radial direction, with blue light mainly absorbed by the outer bark, and far-red-enriched light reaching the underlying xylem and pith. Chlorophylls were evident in the xylem rays and the pith and showed an increasing concentration gradient toward the bark. The stem photosynthetic apparatus showed features typical of acclimation to a low-light environment, such as larger grana stacks, lower chlorophyll a/b and photosystem I/II ratios compared with leaves. Despite likely receiving very few photons, wood chloroplasts were photosynthetically active and fully capable of generating a light-dependent electron transport. Our data provide a comprehensive scenario of the functional features of bark and wood chloroplasts in a woody species and suggest that stem photosynthesis is coherently optimized to the prevailing micro-environmental conditions at the bark and wood level. [ABSTRACT FROM AUTHOR]

Tree breeding programs and wood industries require simple, time- and cost-effective techniques to process large volumes of samples. In recent decades, near-infrared spectroscopy (NIRS) has been acknowledged as one of the most powerful techniques for wood analysis, making it the most used tool for high-throughput phenotyping. Previous studies have shown that a significant number of anatomical, physical, chemical and mechanical wood properties can be estimated through NIRS, both for angiosperm and gymnosperm species. However, the ability of this technique to predict functional traits related to drought resistance has been poorly explored, especially in angiosperm species. This is particularly relevant since determining xylem hydraulic properties by conventional techniques is complex and time-consuming, clearly limiting its use in studies and applications that demand large amounts of samples. In this study, we measured several wood anatomical and hydraulic traits and collected NIR spectra in branches of two Eucalyptus L'Hér species. We developed NIRS calibration models and discussed their ability to accurately predict the studied traits. The models generated allowed us to adequately calibrate the reference traits, with high R 2 (≥0.75) for traits such as P12, P88, the slope of the vulnerability curves to xylem embolism or the fiber wall fraction, and with lower R 2 (0.39–0.52) for P50, maximum hydraulic conductivity or frequency of ray parenchyma. We found that certain wavenumbers improve models' calibration, with those in the range of 4000–5500 cm−1 predicting the highest number of both anatomical and functional traits. We concluded that the use of NIRS allows calibrating models with potential predictive value not only for wood structural and chemical variables but also for anatomical and functional traits related to drought resistance in wood types with complex structure as eucalypts. These results are promising in light of the required knowledge about species and genotypes adaptability to global climatic change. [ABSTRACT FROM AUTHOR]

Wood is a renewable resource that can be employed for the production of second generation biofuels by enzymatic saccharification and subsequent fermentation. Knowledge on how the saccharification potential is affected by genotype-related variation of wood traits and drought is scarce. Here, we used three Populus nigra L. genotypes from habitats differing in water availability to (i) investigate the relationships between wood anatomy, lignin content and saccharification and (ii) identify genes and co-expressed gene clusters related to genotype and drought-induced variation in wood traits and saccharification potential. The three poplar genotypes differed in wood anatomy, lignin content and saccharification potential. Drought resulted in reduced cambial activity, decreased vessel and fiber lumina, and increased the saccharification potential. The saccharification potential was unrelated to lignin content as well as to most wood anatomical traits. RNA sequencing of the developing xylem revealed that 1.5% of the analyzed genes were differentially expressed in response to drought, while 67% differed among the genotypes. Weighted gene correlation network analysis identified modules of co-expressed genes correlated with saccharification potential. These modules were enriched in gene ontology terms related to cell wall polysaccharide biosynthesis and modification and vesicle transport, but not to lignin biosynthesis. Among the most strongly saccharification-correlated genes, those with regulatory functions, especially kinases, were prominent. We further identified transcription factors whose transcript abundances differed among genotypes, and which were co-regulated with genes for biosynthesis and modifications of hemicelluloses and pectin. Overall, our study suggests that the regulation of pectin and hemicellulose metabolism is a promising target for improving wood quality of second generation bioenergy crops. The causal relationship of the identified genes and pathways with saccharification potential needs to be validated in further experiments.

Conifers growing at the alpine timberline are exposed to combinatorial stresses that induce embolism in xylem during winter. We collected branches of Abies mariesii Mast. at the timberline on Mt Norikura of central Japan to evaluate the seasonal changes in the loss of xylem hydraulic conductivity (percent loss of hydraulic conductivity; PLC). Concurrently, we evaluated the distribution of embolized tracheids in native samples via cryo-scanning electron microscopic (cryo-SEM) observation in comparison with the drought-induced embolism samples used for the vulnerability curve. The twigs collected in late winter showed 100 PLC at a water potential of ~−3 MPa, and air-filled tracheids were observed sporadically in the cryo-SEM images. The cryo-SEM images also showed that nearly all pits of the samples from the timberline were aspirated in the xylem with 100 PLC. Conversely, in drought-induced samples used for vulnerability analysis, lower frequency of aspirated pits was observed at similar water potentials and all tracheids in the earlywood of xylem with 100 PLC were filled with air. We hypothesized that pit aspiration is the primary cause of the pronounced winter xylem conductivity loss at the timberline when water potential is near, but still above, the drought-induced vulnerability threshold. Pit aspiration before water loss may be an adaptation to severe winter conditions at timberlines to prevent widespread xylem embolism. The possible causes of pit aspiration are discussed in relation to complex stresses under harsh winter conditions at timberlines. [ABSTRACT FROM AUTHOR]

Sucrose is the central unit of carbon and energy in plants. Active intercellular transport of sucrose is mediated by sucrose transporters (SUTs), genes for which have been found in the genomes of all land plants. However, they have only been assigned functions in angiosperm species. Here, we cloned two types of SUTs from two gymnosperms, the conifers Cedrus deodara (Roxb. G. Don) and Pinus massoniana Lambert, and analyzed their sucrose transport activities. Uptake of the fluorescent sucrose-analog esculin into tobacco epidermis cells expressing the conifer SUT confirmed their transport ability. To determine their function in planta, we investigated their mRNA abundance in relation to photosynthesis and sugar levels in leaves, inner bark, wood and roots. Combined with measurements of protein abundance and immunolocalization of C. deodara SUTs, our results suggest a role for CdSUT1G and CdSUT2 in supporting phloem transport under varying environmental conditions. The implications of these findings regarding conifer physiology and SUT evolution are discussed. [ABSTRACT FROM AUTHOR]

Plants constitute 80% of the biomass on earth, and almost two-thirds of this biomass is found in wood. Wood formation is a carbon (C)-demanding process and relies on C transport from photosynthetic tissues. Thus, understanding the transport process is of major interest for understanding terrestrial biomass formation. Here, we review the molecules and mechanisms used to transport and allocate C in trees. Sucrose is the major form in which C is transported in plants, and it is found in the phloem sap of all tree species investigated so far. However, in several tree species, sucrose is accompanied by other molecules, notably polyols and the raffinose family of oligosaccharides. We describe the molecules that constitute each of these transport groups, and their distribution across different tree species. Furthermore, we detail the metabolic reactions for their synthesis, the mechanisms by which trees load and unload these compounds in and out of the vascular system, and how they are radially transported in the trunk and finally catabolized during wood formation. We also address a particular C recirculation process between phloem and xylem that occurs in trees during the annual cycle of growth and dormancy. A search of possible evolutionary drivers behind the diversity of C-carrying molecules in trees reveals no consistent differences in C transport mechanisms between angiosperm and gymnosperm trees. Furthermore, the distribution of C forms across species suggests that climate-related environmental factors will not explain the diversity of C transport forms. However, the consideration of C-transport mechanisms in relation to tree–rhizosphere coevolution deserves further attention. To conclude the review, we identify possible future lines of research in this field. [ABSTRACT FROM AUTHOR]

Canopy phenology is a widely used proxy for deciduous forest growth with various applications in terrestrial ecosystem modeling. Its use relies on common assumptions that canopy greening and stem growth are tightly coordinated processes, enabling predictions on the timing and the quantity of annual tree growth. Here, we present parallel observations of canopy and stem growth phenology and annual stem increment in around 90 deciduous forest trees with diffuse-porous (Fagus sylvatica, Acer pseudoplatanus, Carpinus betulus) or ring-porous (Quercus robur × petraea) wood anatomy. These data were collected in a mixed temperate forest at the Swiss-Canopy-Crane II site, in 4 years with strongly contrasting weather conditions. We found that stem growth resumption lagged several weeks behind spring canopy greening in diffuse-porous but not in ring-porous trees. Canopy greening and stem growth resumption showed no or only weak signs of temporal coordination across the observation years. Within the assessed species, the seasonal timing of stem growth varied strongly among individuals, as trees with high annual increments resumed growth earlier and also completed their main growth earlier. The length of main growth activity had no influence on annual increments. Our findings not only challenge tight temporal coordination of canopy and stem growth phenology but also demonstrate that longer main growth activity does not translate into higher annual increments. This may compromise approaches modeling tree growth and forest productivity with canopy phenology and growth length., (© The Author(s) 2023. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

The hydraulic death hypothesis suggests that fires kill trees by damaging the plant's hydraulic continuum in addition to stem cambium. A corollary to this hypothesis is that plants that survive fires possess 'pyrohydraulic' traits that prevent heat-induced embolism formation in the xylem and aid post-fire survival. We examine whether hydraulic segmentation within stem xylem may act as such a trait. To do so, we measured the percentage loss of conductance (PLC) and vulnerability to embolism axially along segments of branches exposed to heat plumes in two differing species, fire-tolerant Eucalyptus cladocalyx F. Muell and fire-sensitive Kiggelaria africana L., testing model predictions that fire-tolerant species would exhibit higher degrees of hydraulic segmentation (greater PLC in the distal parts of the branch than the basal) than fire-intolerant species (similar PLC between segments). Following exposure to a heat plume, K. africana suffered between 73 and 84% loss of conductance in all branch segments, whereas E. cladocalyx had 73% loss of conductance in whole branches, including the distal tips, falling to 29% in the most basal part of the branch. There was no evidence for differences in resistance segmentation between the species, and there was limited evidence for differences in distal vulnerability to embolism across the branches. Hydraulic segmentation in E. cladocalyx may enable it to resprout effectively post-fire with a functional hydraulic system. The lack of hydraulic segmentation in K. africana reveals the need to understand possible trade-offs associated with hydraulic segmentation in long-lived woody species with respect to drought and fire., (© The Author(s) 2023. Published by Oxford University Press.)

Since the 1990s the invasive fungus Hymenoscyphus fraxineus has caused severe crown dieback and high mortality rates in Fraxinus excelsior in Europe. In addition to a strong genetic control of tolerance to the fungus, previous studies have found landscape heterogeneity to be an additional driver of variability in the severity of dieback symptoms. However, apart from climatic conditions related to heat and humidity influencing fungal infection success, the mechanistic understanding of why smaller or slower-growing trees are more susceptible to dieback remains less well understood. Here, we analyzed three stands in Switzerland with a unique setting of 8 years of data availability of intra-annual diameter growth and annual crown health assessments. We complemented this by ring width and quantitative wood anatomical measurements extending back before the monitoring started to investigate if wood anatomical adjustments can help better explain the size-related dieback phenomenon. We found that slower-growing trees or trees with smaller crowns already before the arrival of the fungus were more susceptible to dieback and mortality. Defoliation directly reduced growth as well as maximum earlywood vessel size, and the positive relationship between vessel size and growth rate caused a positive feedback amplifying and accelerating crown dieback. Measured non-structural carbohydrate (NSC) concentrations in the outermost five rings did not significantly vary between healthy and weakened trees, which translate into large differences in absolute available amount of NSCs. Thus, we hypothesize that a lack of NSCs (mainly sugars) leads to lower turgor pressure and smaller earlywood vessels in the following year. This might impede efficient water transport and photosynthesis, and be responsible for stronger symptoms of dieback and higher mortality rates in smaller and slower-growing trees. [ABSTRACT FROM AUTHOR]

Compression wood (CW) is a common tissue present in the trunk, branches and roots of mechanically stressed coniferous trees. Its main role is to increase the mechanical strength and regain the vertical orientation of a leaning stem. Compression wood is thought to influence the climate signal in different tree-ring measures. Hence trees containing CW are mostly excluded from tree-ring studies reconstructing past climate variability. There is a large gap of systematic work testing the potential effect of CW on the strength of the climate signal in different tree-ring parameters, especially stable isotope records. Here we test for the first time the effect of CW contained in montane Norway spruce (Picea abies L. Karst) on both δ13C and δ18O tree-ring cellulose records by analyzing compression and opposite wood radii from several disturbed trees together with samples from undisturbed reference trees. We selected four trees tilted by geomorphic processes that were felled by wind and four undisturbed reference trees in the Tatra Mountains, Poland. We qualitatively classified the strength of CW using wood cell anatomical characteristics (tracheid shape, cell wall thickness and presence of intercellular spaces). Then we developed tree-ring width and δ13C and δ18O chronologies from the CW radii, from the opposite radii of the tilted trees and from the reference radii. We tested the effect of CW on tree-ring cellulose δ13C and δ18O variability and on the climate signal strength. We found only minor differences in the means of δ13C and δ18O of compression (δ13C: −22.81‰, δ18O: 28.29‰), opposite (δ13C: −23.02‰; δ18O: 28.05‰) and reference (δ13C: −22.78‰; δ18O: 27.61‰) radii. The statistical relationships between climate variables, δ13C and δ18O, remained consistent among all chronologies. Our findings suggest that moderately tilted trees containing CW can be used to reconstruct past geomorphic activity and for stable isotope-based dendroclimatology. [ABSTRACT FROM AUTHOR]

Gas inside wood plays an important role in plant functioning, but there has been no study examining the adaptive nature of gas inside wood across plants differing in biomechanical demands. Using a comparative approach, we measured gas volumetric content, xylem's anatomical traits and wood density of 15 tree and 16 liana species, to test whether gas content varies between these plant types strongly differing in their biomechanical demands. We asked (i) whether trees and lianas differ in gas content and (ii) how anatomical traits and wood density are related to gas content. Lianas had significantly less gas content in their branches compared with tree species. In tree species, gas content scaled positively with fiber, vessel and xylem cross-sectional area and fiber and vessel diameter, and negatively with dry-mass density. When pooling trees and lianas together, fiber cross-sectional area was the strongest predictor of gas content, with higher xylem cross-sectional area of fiber associated with higher gas content. In addition, we showed, through a simple analytical model, that gas inside wood increases the minimum branch diameter needed to prevent rupture, and this effect was stronger on trees compared with lianas. Our results support the view that gas inside wood plays an important role in the evolution of biomechanical functioning in different plant forms. Gas inside wood may also play an important role in physiological activities such as water transport, storage, photosynthesis and respiration, but it is still unknown whether these roles are or are not secondary to the mechanical support. [ABSTRACT FROM AUTHOR]

Cork cambium (or phellogen) is a secondary meristem responsible for the formation of phelloderm and phellem/cork, which together compose the periderm. In Quercus suber L. , the phellogen is active throughout the entire life of the tree, producing a continuous and renewable outer bark of cork. To identify specific candidate genes associated with cork cambium activity and phellem differentiation, we performed a comparative transcriptomic study of Q. suber secondary growth tissues (xylem and phellogen/phellem) using RNA-seq. The present work provides a high-resolution map of all the transcripts identified in the phellogen/phellem tissues. A total of 6013 differentially expressed genes were identified, with 2875 of the transcripts being specifically enriched during the cork formation process versus secondary xylem formation. Furthermore, cork samples originating from the original phellogen ('virgin' cork) and from a traumatic phellogen ('amadia' cork) were also compared. Our results point to a shortlist of potentially relevant candidate genes regulating phellogen activity and phellem differentiation, including novel genes involved in the suberization process, as well as genes associated to ethylene and jasmonate signaling and to meristem function. The future functional characterization of some of the identified candidate genes will help to elucidate the molecular mechanisms underlying cork cambium activity and phellem differentiation. [ABSTRACT FROM AUTHOR]

The regulation of DNA accessibility by histone modification has emerged as a paradigm of developmental and environmental programming. Chromatin immunoprecipitation followed by sequencing (ChIP-seq) is a versatile tool to investigate in vivo protein-DNA interaction and has enabled advances in mechanistic understanding of physiologies. The technique has been successfully demonstrated in several plant species and tissues; however, it has remained challenging in woody tissues, in particular complex structures such as perennating buds. Here we developed a ChIP method specifically for mature dormant buds of grapevine (Vitis vinifera cv. Cabernet Sauvignon). Each step of the protocol was systematically optimized, including crosslinking, chromatin extraction, sonication and antibody validation. Analysis of histone H3-enriched DNA was performed to evaluate the success of the protocol and identify occupancy of histone H3 along grapevine bud chromatin. To our best knowledge, this is the first ChIP experiment protocol optimized for the grapevine bud system., (© The Author(s) 2023. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Xylem anatomy may change in response to environmental or biotic stresses. Vascular occlusion, an anatomical modification of mature xylem, contributes to plant resistance and susceptibility to different stresses. In woody organs, xylem occlusions have been examined as part of the senescence process, but their presence and function in leaves remain obscure. In grapevine, many stresses are associated with premature leaf senescence inducing discolorations and scorched tissue in leaves. However, we still do not know whether the leaf senescence process follows the same sequence of physiological events and whether leaf xylem anatomy is affected in similar ways. In this study, we quantified vascular occlusions in midribs from leaves with symptoms of the grapevine disease esca, magnesium deficiency and autumn senescence. We found higher amounts of vascular occlusions in leaves with esca symptoms (in 27% of xylem vessels on average), whereas the leaves with other symptoms (as well as the asymptomatic controls) had far fewer occlusions (in 3% of vessels). Therefore, we assessed the relationship between xylem occlusions and esca leaf symptoms in four different countries (California in the USA, France, Italy and Spain) and eight different cultivars. We monitored the plants over the course of the growing season, confirming that vascular occlusions do not evolve with symptom age. Finally, we investigated the hydraulic integrity of leaf xylem vessels by optical visualization of embolism propagation during dehydration. We found that the occlusions lead to hydraulic dysfunction mainly in the peripheral veins compared with the midribs in esca symptomatic leaves. These results open new perspectives on the role of vascular occlusions during the leaf senescence process, highlighting the uniqueness of esca leaf symptoms and its consequence on leaf physiology., (© The Author(s) 2022. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

The drought susceptibility of woody saplings may explain their low survival in arid environments. Therefore, it is critical to determine which morphological and physiological traits are more responsive to drought among young plants. This study tested whether plant responses to experimental drought differ between two plant functional groups: the deciduous and evergreen species. We predicted that deciduous species would present a tighter stomatal control under drought, coupled with fast carbon fixation under no stress, tending toward isohydry and faster growth rates than the evergreen species. Using 1-year-old saplings from three evergreen and four deciduous Sonoran Desert tree species, we evaluated their hydraulic and gas exchange traits under three experimental irrigation conditions: high, intermediate and low water availability. We measured CO2 assimilation rates (A), stomatal conductance (gs), the level of iso-anisohydry (as the plant's ability to maintain constant their water potential) and seven morphological and growth-related traits throughout 2 months. Under high water availability, saplings reached their maximum values of A and gs, which were significantly higher for deciduous than evergreen species. Correlations among hydroscape area (HA) and leaf traits positioned species along the iso/anisohydric continuum. Deciduous species presented isohydric characteristics, including low HA, high gs, A and Huber values (HVs), and traits indicative of a faster use of resources, such as low stem-specific density (SSD) and low leaf mass per area (LMA). By contrast, evergreen species showed traits that indicate slow resource use and anisohydric behavior, such as high HA, SSD and LMA, and low gs, A and HVs. Deciduous species drastically reduced gas exchange rates in response to drought, while evergreen maintained low rates independently of drought intensity. Overall, desert saplings showed strategies concordant with the iso-anisohydric continuum and the fast-slow use of resources., (© The Author(s) 2022. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Climate change is increasing the severity and duration of drought events experienced by forest ecosystems. Because water is essential for tree physiological processes, the ability of trees to survive prolonged droughts will largely depend on whether they have access to reliable water sources. While many woody plant species exhibit the ability to shift water sources between different depths of soil and rock water in response to changes in climate and water availability, it is unclear if Sierra Nevada conifers exhibit this plasticity. Here we analysed the δ18O and δ13C values of annual tree rings to determine the water-use patterns of large Sierra Nevada conifers during the 2012-16 California drought and 4 years before this drought event (2004-07). We analysed four species (Pinus jeffreyi Grev. & Balf. (Jeffrey pine), Pinus lambertiana Dougl. (sugar pine), Abies concolor (Gord. & Glend.) Lindl. Ex Hilderbr (white fir) and Calocedrus decurrens (Torr.) Florin (incense-cedar)) across a range of topographic positions to investigate differences in water-use patterns by species and position on the landscape. We found no significant differences in δ18O and δ13C values for the pre-drought and drought periods. This stability in δ18O values suggests that trees did not shift their water-use patterns in response to the 2012-16 drought. We did find species-specific differences in water-use patterns, with incense-cedar exhibiting more depleted δ18O values than all other species. We also found trends that suggest the water source used by a tree may depend on topographic and growing environment attributes such as topographic wetness and the surrounding basal area. Overall, our results suggest that the water source used by trees varies by the species and topographic position, but that Sierra Nevada conifers do not switch their water-use patterns in response to the drought. This lack of plasticity could make Sierra Nevada conifers particularly vulnerable to drought mortality as their historically reliable water sources begin to dry out with climate change., (© The Author(s) 2022. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

The hydraulic traits of a plant species may reflect its climate adaptations. Southwest China is considered as a biodiversity hotpot of the genus Quercus (oak). However, the hydraulic adaptations of Asian oaks to their climate niches remain unclear. Ten common garden-grown oak species with distinct natural distributions in eastern Asia were used to determine their stem xylem embolism resistance (water potential at 50% loss of hydraulic conductivity, P50), stem hydraulic efficiency (vessel anatomy and sapwood specific hydraulic conductivity (Ks)) and leaf anatomical traits. We also compiled four key functional traits: wood density, hydraulic-weighted vessel diameter, Ks and P50 data for 31 oak species from previous literature. We analyzed the relationship between hydraulic traits and climatic factors over the native ranges of 41 oak species. Our results revealed that the 10 Asian oak species, which are mainly distributed in humid subtropical habitats, possessed a stem xylem with low embolism resistance and moderate hydraulic efficiency. The deciduous and evergreen species of the 10 Asian oaks differed in the stem and leaf traits related to hydraulic efficiency. Ks differed significantly between the two phenological groups (deciduous and evergreens) in the 41-oak dataset. No significant difference in P50 between the two groups was found for the 10 Asian oaks or the 41-oak dataset. The oak species that can distribute in arid habitats possessed a stem xylem with high embolism resistance. Ks negatively related to the humidity of the native range of the 10 Asian oaks, but showed no trend when assessing the entire global oak dataset. Our study suggests that stem hydraulic conductivity and embolism resistance in Quercus species are shaped by their climate niche. Our findings assist predictions of oak drought resistance with future climate changes for oak forest management., (© The Author(s) 2022. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Carbon fractions of tree tissues are a key component of forest carbon mass estimation. Several methods have been used to measure carbon fractions, yet no comprehensive comparison between methods has been performed. We found significant differences between carbon fractions derived from four sample preparation methods: oven-drying, vacuum desiccation, freeze-drying, and a new method that consisted of (i) not drying samples, (ii) cutting samples instead of grinding them, (iii) measuring carbon content of samples, (iv) oven-drying remaining sample material and (v) using mass measurements of remaining sample material before and after oven-drying to adjust measured carbon fraction values to an oven-dry basis (minimize the loss of carbon (MLC) method). Oven-drying, freeze-drying and vacuum desiccation resulted in lower average carbon fraction estimates than the MLC method, suggesting that they do not capture as much of the carbon present in tree tissues. Further analysis showed significant, though small, differences in carbon fractions between powdered samples and samples excised from tree core segments with a razor blade. Powdered samples were found to have lower carbon fractions than the excised samples, indicating that some carbon is lost when samples are powdered instead of cut. Utilization of the MLC method captured an average of 1.4% more carbon on an oven-drying basis than freeze-drying, the next best method. Additionally, when applied to different tree tissue types, these methods measured different volatile carbon fractions, indicating that studies attempting to quantify volatile carbon and total carbon fraction in trees should measure all tissue types present. [ABSTRACT FROM AUTHOR]

Seasonal analyses of cambial cell production and day-by-day stem radial increment can help to elucidate how climate modulates wood formation in conifers. Intra-annual dynamics of wood formation were determined with microcores and dendrometers and related to climatic signals in Norway spruce (Picea abies (L.) Karst.). The seasonal dynamics of these processes were observed at two sites of different altitude, Savignano (650 m a.s.l.) and Lavazè (1800 m a.s.l.) in the Italian Alps. Seasonal dynamics of cambial activity were found to be site specific, indicating that the phenology of cambial cell production is highly variable and plastic with altitude. There was a site-specific trend in the number of cells in the wall thickening phase, with the maximum cell production in early July (DOY 186) at Savignano and in mid-July (DOY 200) at Lavazè. The formation of mature cells showed similar trends at the two sites, although different numbers of cells and timing of cell differentiation were visible in the model shapes; at the end of ring formation in 2010, the number of cells was four times higher at Savignano (106.5 cells) than at Lavazè (26.5 cells). At low altitudes, microcores and dendrometers described the radial growth patterns comparably, though the dendrometer function underlined the higher upper asymptote of maximum growth in comparison with the cell production function. In contrast, at high altitude, these functions exhibited different trends. The best model was obtained by fitting functions of the Gompertz model to the experimental data. By combining radial growth and cambial activity indices we defined a model system able to synchronize these processes. Processes of adaptation of the pattern of xylogenesis occurred, enabling P. abies to occupy sites with contrasting climatic conditions. The use of daily climatic variables in combination with plant functional traits obtained by sensors and/or destructive sampling could provide a suitable tool to better investigate the effect of disturbances on response strategies in trees and, consequently, contribute to improving our prediction of tree growth and species resilience based on climate scenarios. [ABSTRACT FROM AUTHOR]

Leaf trait variation enables plants to utilize large gradients of light availability that exist across canopies of high leaf area index (LAI), allowing for greater net carbon gain while reducing light availability for understory competitors. While these canopy dynamics are well understood in forest ecosystems, studies of canopy structure of woody shrubs in grasslands are lacking. To evaluate the investment strategy used by these shrubs, we investigated the vertical distribution of leaf traits and physiology across canopies of Cornus drummondii, the predominant woody encroaching shrub in the Kansas tallgrass prairie. We also examined the impact of disturbance by browsing and grazing on these factors. Our results reveal that leaf mass per area (LMA) and leaf nitrogen per area (Na) varied approximately threefold across canopies of C. drummondii, resulting in major differences in the physiological functioning of leaves. High LMA leaves had high photosynthetic capacity, while low LMA leaves had a novel strategy for maintaining light compensation points below ambient light levels. The vertical allocation of leaf traits in C. drummondii canopies was also modified in response to browsing, which increased light availability at deeper canopy depths. As a result, LMA and Na increased at lower canopy depths, leading to a greater photosynthetic capacity deeper in browsed canopies compared to control canopies. This response, along with increased light availability, facilitated greater photosynthesis and resource-use efficiency deeper in browsed canopies compared to control canopies. Our results illustrate how C. drummondii facilitates high LAI canopies and a compensatory growth response to browsing-both of which are key factors contributing to the success of C. drummondii and other species responsible for grassland woody encroachment., (© The Author(s) 2022. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Estimates of tree heartwood and sapwood profiles are important in the pulp industry and for dynamic vegetation models, in which they determine tree biomechanical stability and hydraulic conductivity. Several phenomenological models of stem profiles have been developed for this purpose, based on assumptions on how tree crown and foliage distributions change over time. Here, we derive estimates of tree profiles by synthesizing a simple pipe model theory of plant form with a recently developed theory of branch thinning that from simple assumptions quantifies discarded branches and leaves. This allows us to develop a new trunk model of tree profiles from breast height up to the top of the tree. We postulate that leaves that are currently on the tree are connected by sapwood pipes, while pipes that previously connected discarded leaves or branches form the heartwood. By assuming that a fixed fraction of all pipes remain on the trunk after a branching event, as the trunk is traversed from the root system to the tips, this allows us to quantify trunk heartwood and sapwood profiles. We test the trunk model performance on empirical data from five tree species across three continents. We find that the trunk model accurately describes heartwood and sapwood profiles of all tested tree species (calibration; R2: 84-99%). Furthermore, once calibrated to a tree species, the trunk model predicts heartwood and sapwood profiles of conspecific trees in similar growing environments based only on the age and height of a tree (cross-validation/prediction; R2: 68-98%). The fewer and often contrasting parameters needed for the trunk model make it a potentially useful complementary tool for biologists and foresters., (© The Author(s) 2022. Published by Oxford University Press.)

An increase in temperature along with a decrease in summer precipitation in Central Europe will result in an increased frequency of drought events and gradually lead to a change in species composition in forest ecosystems. In the present study, young oaks (Quercus robur L. and Quercus petraea (Matt.) Liebl.) were transplanted into large mesocosms and exposed for 3 years to experimental warming and a drought treatment with yearly increasing intensities. Carbon and oxygen isotopic (δ13C and δ18O) patterns were analysed in leaf tissue and tree-ring cellulose and linked to leaf physiological measures and tree-ring growth. Warming had no effect on the isotopic patterns in leaves and tree rings, while drought increased δ18O and δ13C. Under severe drought, an unexpected isotopic pattern, with a decrease in δ18O, was observed in tree rings but not in leaves. This decrease in δ18O could not be explained by concurrent physiological analyses and is not supported by current physiological knowledge. Analysis of intra-annual tree-ring growth revealed a drought-induced growth cessation that interfered with the record of isotopic signals imprinted on recently formed leaf carbohydrates. This missing record indicates isotopic uncoupling of leaves and tree rings, which may have serious implications for the interpretation of tree-ring isotopes, particularly from trees that experienced growth-limiting stresses. [ABSTRACT FROM AUTHOR]