Quantitative Wood Anatomy Research Articles

Revealing Polylepis microphylla as a suitable tree species for dendrochronology and quantitative wood anatomy in the Andean montane forests

In the tropical Andes climate change is expected to increase temperatures and change precipitation patterns. To overcome the lack of systematic weather records that limits the performance of climate models in this region, the use of the environmental information contained in tree rings from tropical Andean species have been found useful to reconstruct spatio-temporal climate variability. Because classical dendrochronology based on ring-width patterns is often challenging in the tropics, alternative approaches such as Quantitative Wood Anatomy (QWA) based on the measurement and quantification of anatomical traits within tree rings can be a significant advance in the field. Here we assess the dendrochronological potential of Polylepis microphylla and its climate sensitivity by using i) classic dendrochronological methods to generate the first Tree-ring Width (TRW) chronology for this tree species spanning from 1965 to 2018; ii) radiocarbon (¹⁴C) analyses as an independent validation method to assess the annual periodicity of the tree growth layers; and iii) QWA to generate tree-ring annual records of the number (VN) and size (VS) of vessels to investigate the climate sensitivity of these anatomical traits. The annual periodicity in P. microphylla radial growth was confirmed by both dendrochronological and ¹⁴C analyses. We found that VN and VS are promising new proxies to reconstruct climate variability in this region and that they provide different information than TRW. While TRW provides information at inter-annual resolution (i.e., year-to-year variability), VN and VS generated with sectorial QWA provide intra-annual resolution for each stage of the growing process. The TRW and the anatomical traits (i.e., VN and VS) showed strong positive correlation with maximum temperature for different periods of the growing season: while VS is higher with warmer conditions prior to the growing season onset, tree-rings are wider and present higher number of vessels when warmer conditions occur during the current growing season. Our findings pointed out the suitability of P. microphylla for dendrochronological studies and may suggest a good performance of this species under the significant warming expected according to future projections for the tropical Andes.

Characteristics of a multi-species conifer network of wood properties chronologies from Southern Australia

Progress in quantitative wood anatomy has resulted in a growing number of increasingly understood proxies from the tree-ring archive. Much of this work has been based on tree species in the Northern Hemisphere. Here, we present and examine a relatively dense network of wood property chronologies (wood density, tracheid radial diameter, cell wall thickness and ring width) from several species in Tasmania, southern Australia. We ask how the relationships amongst the different types of chronologies differ within and amongst species. We also consider how each chronology responds to monthly climate. In general terms, and similar to findings in the Northern Hemisphere, relationships between the various wood properties and climate are stronger than those between climate and ring width chronologies. An important exception to this is the highest elevation Lagarostrobos franklinii site. Additionally, strongest response to climate for the wood properties generally occurs for the concurrent growing season compared to the prior growing season for ring width. Relationships amongst the various chronology types differ for the various species, with L. franklinii also showing some variation in these relationships by site (possibly associated with elevation). Results suggest there is considerable value in further exploring the potential for developing anatomical wood chronologies for climate reconstruction from other species for which ring widths do not exhibit a strong climate signal.

Can quantitative wood anatomy data coupled with machine learning analysis discriminate CITES species from their look-alikes?

Can quantitative wood anatomy data coupled with machine learning analysis discriminate CITES species from their look-alikes?

Tracheidogram’s Classification as a New Potential Proxy in High-Resolution Dendroclimatic Reconstructions

Quantitative wood anatomy (QWA) is widely used to resolve a fundamental problem of tree responses to past, ongoing and forecasted climate changes. Potentially, QWA data can be considered as a new proxy source for long-term climate reconstruction with higher temporal resolution than traditional dendroclimatic data. In this paper, we considered a tracheidogram as a set of two interconnected variables describing the dynamics of seasonal variability in the radial cell size and cell wall thickness in conifer trees. We used 1386 cell profiles (tracheidograms) obtained for seven Scots pine (Pinus sylvestris) trees growing in the cold semiarid conditions of Southern Siberia over the years 1813–2018. We developed a “deviation tracheidogram” approach for adequately describing the traits of tree-ring formation in different climate conditions over a long-term time span. Based on the NbClust approach and K-means method, the deviation tracheidograms were reliably split into four clusters (classes) with clear bio-ecological interpretations (from the most favorable growth conditions to worse ones) over the years 1813–2018. It has been shown that the obtained classes of tracheidograms can be directly associated with different levels of water deficit, for both the current and previous growing seasons. The tracheidogram cluster reconstruction shows that the entire 19th century was characterized by considerable water deficit, which has not been revealed by the climate-sensitive tree-ring chronology of the study site. Therefore, the proposed research offers new perspectives for better understanding how tree radial growth responds to changing seasonal climate and a new independent proxy for developing long-term detailed climatic reconstructions through the detailed analysis of long-term archives of QWA data for different conifer species and various forest ecosystems in future research.

Drought impacts on tree carbon sequestration and water use - evidence from intra-annual tree-ring characteristics.

Summary The impact of climate extremes on forest ecosystems is poorly understood but important for predicting carbon and water cycle feedbacks to climate. Some knowledge gaps still remain regarding how drought‐related adjustments in intra‐annual tree‐ring characteristics directly impact tree carbon and water use.In this study we quantified the impact of an extreme summer drought on the water‐use efficiency and carbon sequestration of four mature Norway spruce trees. We used detailed observations of wood formation (xylogenesis) and intra‐annual tree‐ring properties (quantitative wood anatomy and stable carbon isotopes) combined with physiological water‐stress monitoring.During 41 d of tree water deficit, we observed an enrichment in 13C but a reduction in cell enlargement and wall‐thickening processes, which impacted the anatomical characteristics. These adjustments diminished carbon sequestration by 67% despite an 11% increase in water‐use efficiency during drought. However, with the resumption of a positive hydric state in the stem, we observed a fast recovery of cell formation rates based on the accumulated assimilates produced during drought.Our findings enhance our understanding of carbon and water fluxes between the atmosphere and forest ecosystems, providing observational evidence on the tree intra‐annual carbon sequestration and water‐use efficiency dynamics to improve future generations of vegetation models.

Relationships Between Wood-Anatomical Features and Resistance Drilling Density in Norway Spruce and European Beech

Environmental conditions affect tree-ring width (TRW), wood structure, and, consequently, wood density, which is one of the main wood quality indicators. Although studies on inter- and intra-annual variability in tree-ring features or density exist, studies demonstrating a clear link between wood structure on a cellular level and its effect on wood density on a macroscopic level are rare. Norway spruce with its simple coniferous structure and European beech, a diffuse-porous angiosperm species were selected to analyze these relationships. Increment cores were collected from both species at four sites in Slovenia. In total, 24 European beech and 17 Norway spruce trees were sampled. In addition, resistance drilling measurements were performed just a few centimeters above the increment core sampling. TRW and quantitative wood anatomy measurements were performed on the collected cores. Resistance drilling density values, tree-ring (TRW, earlywood width–EWW, transition-TWW, and latewood width–LWW) and wood-anatomical features (vessel/tracheid area and diameter, cell density, relative conductive area, and cell wall thickness) were then averaged for the first 7 cm of measurements. We observed significant relationships between tree-ring and wood-anatomical features in both spruce and beech. In spruce, the highest correlation values were found between TRW and LWW. In beech, the highest correlations were observed between TRW and cell density. There were no significant relationships between wood-anatomical features and resistance drilling density in beech. However, in spruce, a significant negative correlation was found between resistance drilling density and tangential tracheid diameter, and a positive correlation between resistance drilling density and both TWW + LWW and LWW. Our findings suggest that resistance drilling measurements can be used to evaluate differences in density within and between species, but they should be improved in resolution to be able to detect changes in wood anatomy.

Dieter Eckstein – A cornerstone of European dendrochronology

On November 10th 2021, Dieter Eckstein passed away at age 82. Born and raised as a forester’s child, his entire life was connected to trees and wood. He grew up to become a dedicated scientist and teacher. His legacy includes both his own considerable research accomplishments as well as his founding of a growing network of tree biologists and wood scientists. From his doctoral degree onwards, the concepts and applications of dendrochronology were his passion, motivated by great curiosity in environmental influences on tree growth. He proved that dendroarchaeology can be accurate and precise, even for timber grown in the mild European maritime climate. He pioneered both techniques and concepts of xylogenesis and quantitative wood anatomy and advanced the potential for tropical dendrochronology. In all of these accomplishments, Dieter collaborated with students and colleagues from all over the world. His Dendrochronological Laboratory at the University of Hamburg hosted both young and experienced scientists from many countries. The European Working Group on Dendrochronology, which he founded in the early 1990s, was his natural habitat and playground to invent and present new research activities. We and the entire dendrochronology community have lost an inspiring colleague and visionary.

Landslide-induced changes in tree-ring anatomy: A new dendrogeomorphic avenue?

The dendrogeomorphic dating of past landslides can be limited by the speed or magnitude of movements, as they are sometimes too small to induce growth responses in trees standing on unstable slopes. As a result, movements of low magnitude are generally assumed to remain undetectable in standard macroscopic analysis of growth disturbances in tree-ring series. To close this gap and to enlarge the suite of approaches available for the dendrogeomorphic detection of landslide activity in growth-ring records of trees, this study focuses on the detection of wood-anatomical growth responses in trees following low-magnitude landslides.Anatomical growth responses of five individuals of common spruce (Picea abies (L.) Karst.) growing on a monitored landslide have been analyzed in this study on a total of 200 tree rings from 40 increment cores extracted at two vertical levels of tree stems. Analysis included seven anatomical parameters commonly used in quantitative wood anatomy, with a focus on the year 2017 year during which the landslide body moved by 8 mm (prior to the start of the growing season) according to extensometer records. We report that mean tracheid lumen area (TLA) decreased significantly due to landsliding, whereas the mean number of traumatic resin ducts (TRD) and rays increased significantly after the occurrence of the landslide. By contrast, the intensity of stem tilting was visibly too small to induce the formation of compression wood but strong enough to induce larger number of rays, which are known to enhance stability as well. We also realize that the low intensity of the landslide movement also resulted in rather localized anatomical anomalies along the stem, with detected responses decreasing in intensity with increasing distance from the ground. We conclude that anatomical responses of P. abies trees can be used to detect past activity in low-magnitude landsliding and that this new proxy can thus extend the applicability of tree-ring approaches to landslide research and to the detection of events that have hitherto remained unidentified in reconstructions.

495-Year Wood Anatomical Record of Siberian Stone Pine (Pinus sibirica Du Tour) as Climatic Proxy on the Timberline

The application of quantitative wood anatomy (QWA) in dendroclimatic analysis offers deep insight into the climatic effect on tree-ring formation, which is crucial in understanding the forests’ response to climate change. However, interrelations between tree-ring traits should be accounted to separate climatic signals recorded during subsequent stages of cell differentiation. The study was conducted in the South Siberian alpine timberline on Pinus sibirica Du Tour, a species considered unpromising in dendroclimatology. Relationships between tree-ring width, cell number N, mean and maximum values of radial diameter D, and cell wall thickness (CWT) were quantified to obtain indexed anatomical chronologies. Exponential functions with saturation D(N) and CWT(N) were proposed, which explained 14–69% and 3–61% of their variability, respectively. Indexation unabated significance of the climatic signals but separated them within a season. Analysis of pointer years and climatic extremes revealed predominantly long-term climatogenic changes of P. sibirica radial growth and QWA and allowed to obtain QWA-based 11-year filtered reconstructions of vegetative season climatic characteristics (R2adj = 0.32–0.66). The revealed prevalence of low-frequency climatic reactions is probably explained by a strategy of slow accumulation and utilization of resources implemented by P. sibirica. It makes this species’ QWA a promising proxy for decadal climatic variations in various intra-seasonal timeframes.

The Origin of Tree‐Ring Reconstructed Summer Cooling in Northern Europe During the 18th Century Eruption of Laki

AbstractBasaltic fissure eruptions, which are characteristic of Icelandic volcanism, are extremely hazardous due to the large quantities of gases and aerosols they release into the atmosphere. The 1783–1784 CE Laki eruption was one of the most significant high‐latitude eruptions in the last millennium and had substantial environmental and climatic impacts. Contemporary observations recorded the presence of a sulfuric haze over Iceland and Europe, which caused famine from vegetation damage and resulted in a high occurrence of respiratory illnesses and related mortality. Historical records in northern Europe show that the summer of 1783 was anomalously warm, but regional tree‐ring maximum latewood density (MXD) data from that year are low and lead to erroneously colder reconstructed summer temperatures. Here, we measure wood anatomical characteristics of Scots pine (Pinus sylvestris) from Jämtland, Sweden in order to identify the cause of this discrepancy. We show that the presence of intraannual density fluctuations in the majority of 1783 growth rings, a sudden reduction in lumen and cell wall area, and the measurement resolution of traditional X‐ray densitometry led to the observed reduced annual MXD value. Multiple independent lines of evidence suggest these anatomical anomalies were most likely the result of direct acidic damage to trees in Northern Europe and that the normal relationship between summer temperature and MXD can be disrupted by this damage. Our study also demonstrates that quantitative wood anatomy offers a high‐resolution approach to identifying anomalous years and extreme events in the tree‐ring record.

Mask, Train, Repeat! Artificial Intelligence for Quantitative Wood Anatomy.

The recent developments in artificial intelligence have the potential to facilitate new research methods in ecology. Especially Deep Convolutional Neural Networks (DCNNs) have been shown to outperform other approaches in automatic image analyses. Here we apply a DCNN to facilitate quantitative wood anatomical (QWA) analyses, where the main challenges reside in the detection of a high number of cells, in the intrinsic variability of wood anatomical features, and in the sample quality. To properly classify and interpret features within the images, DCNNs need to undergo a training stage. We performed the training with images from transversal wood anatomical sections, together with manually created optimal outputs of the target cell areas. The target species included an example for the most common wood anatomical structures: four conifer species; a diffuse-porous species, black alder (Alnus glutinosa L.); a diffuse to semi-diffuse-porous species, European beech (Fagus sylvatica L.); and a ring-porous species, sessile oak (Quercus petraea Liebl.). The DCNN was created in Python with Pytorch, and relies on a Mask-RCNN architecture. The developed algorithm detects and segments cells, and provides information on the measurement accuracy. To evaluate the performance of this tool we compared our Mask-RCNN outputs with U-Net, a model architecture employed in a similar study, and with ROXAS, a program based on traditional image analysis techniques. First, we evaluated how many target cells were correctly recognized. Next, we assessed the cell measurement accuracy by evaluating the number of pixels that were correctly assigned to each target cell. Overall, the “learning process” defining artificial intelligence plays a key role in overcoming the issues that are usually manually solved in QWA analyses. Mask-RCNN is the model that better detects which are the features characterizing a target cell when these issues occur. In general, U-Net did not attain the other algorithms’ performance, while ROXAS performed best for conifers, and Mask-RCNN showed the highest accuracy in detecting target cells and segmenting lumen areas of angiosperms. Our research demonstrates that future software tools for QWA analyses would greatly benefit from using DCNNs, saving time during the analysis phase, and providing a flexible approach that allows model retraining.

Anatomical comparison of branches and trunks of seven commercial wood species

Indonesia is a mega-biodiversity country that grows about 4000 timber producer tree species in tropical rain forest. The comprehensive information of wood properties and quality is important in managing the natural resources sustainably. However, stem in basic properties studies of wood from the natural forest is limited because of some difficulties in harvesting until the transportation process. Hence, study the wood branches becomes a solution, as sometimes wood core samples are not adequate. The question was whether the branch properties could represent the main stem properties? Wood anatomy is an important wood property that can predict the other properties, for instance, the physical and mechanical properties, that determine the effective use of this material. This paper aims to present the comparison of wood anatomical properties of branches and main stem of seven commercial wood species. Quantitative and qualitative anatomical structures were investigated according to the International Association of Wood Anatomist (IAWA) Committee. Result shows that the quantitative wood anatomy of four samples (Mimba, Leda, Jabon, and Bintangur) was statistically different, while only one parameter in Tusam, Mindi, and Khaya was different. Accordingly, it can be concluded that studying the wood properties could be carried out using branch effectively.

Q-NET – a new scholarly network on quantitative wood anatomy

Quantitative wood anatomy (QWA) is a dynamic research approach of increasing interest that can provide answers to a wide range of research questions across different disciplines. However, the lack of common protocols and knowledge gaps hinder the realisation of the full potential of QWA. Therefore, we established the new community-based network Q-NET to provide an open interdisciplinary platform for exchange and research around QWA. Q-NET (https://qwa-net.com) combines an online knowledge and exchange base with virtual workshops. The first two workshops each attracted more than 125 participants from around the world, demonstrating the community's interest in QWA and this virtual way of networking and collaborating. Indeed, virtual networks such as Q-NET could increase the inclusiveness, efficiency and sustainability of scientific collaboration while providing additional training and teaching opportunities for early career scientists, both of which complement in-person conferences and workshops.

Warming menaces high-altitude Himalayan birch forests: Evidence from cambial phenology and wood anatomy

The Himalayas are experiencing dramatic warming and drought events, which occur at a faster rate than the global average. How will the high-elevation ecosystems cope with such changing conditions? This study aims to combine cambial phenology with quantitative wood anatomy to understand how the growth dynamics affect the trade-off between safety and efficiency of water transport in the Himalayan forests. We investigated the timing of xylogenesis and its relationship with hydraulic wood anatomical traits in Himalayan birch (Betula utilis D. Don) at two altitudes in the central Himalayas. Xylogenesis was monitored weekly from April to September 2014 by collecting microcores. We measured vessel area on histological sections, calculated the hydraulic diameter, and assessed the effects of climate on xylem production rate. Xylogenesis occurred from June to September in both study sites. Moisture availability influenced positively xylem growth besides thermal conditions. The onset and duration of vessel enlargement were correlated with the hydraulic diameter. Trees with earlier onsets and longer durations of vessel enlargement exhibited larger hydraulic diameters. The close relationship between cambial phenology and hydraulic diameter suggests that earlier growth resumption induced by climate changes may result in larger vessels, which are more efficient in water transport, but vulnerable to drought-induced cavitation. The ongoing warming and drying climate conditions may menace the survival of birch forest in the central Himalayas.

Xylem features detrending methods matter: A case study on earlywood vessels of Fraxinus mandshurica

Xylem anatomy can provide valuable information about tree allometry and ecophysiological performance. It has unique advantages in the study of global climate change and forest adaptation compared to traditional ring-width or density proxy parameters. Developing appropriate detrended sequences of xylem features is necessary to carry out a series of further studies. However, the differences between detrending methods and their effects for xylem features are still unclear, presenting a challenge to the choice of proper detrending methods. In this study, we used the earlywood vessel features of Fraxinus mandshurica as an example and compared the differences in four common detrending methods (Spline, CSS: cubic smoothing spline, NEP: negative exponential curve, and RCS: regional curve standardization) to highlight the importance of the detrending methods in dendroanatomical studies. The results show that detrending methods significantly affect the long-term growth trends and climate signals recorded in xylem anatomical features. Among them, NEP has good performance in terms of ring width and vessel number, which is not suitable for other vessel features because of its deterministic nature. Spline and CSS are relatively conservative methods, in which are not easy to make mistakes, but the problem of overfitting (lack of climate signals) deserves attention. Multiple RCS may be a promising detrending method with quantitative wood anatomy technology development if sufficient samples are available. Our results highlight the importance of selecting detrending methods for xylem feature identification and emphasize the importance of testing various detrending methods rather than arbitrarily choosing a standard method. The detrending method should be selected based on the purpose, such as the balance between climate signals and thorough detrended effects. We recommend applying multiple methods to test the growth trends in xylem features before selecting a detrending method. More interdisciplinary studies involving ecophysiology, wood anatomy, and dendrochronology will help to distil the valuable information in xylem features accurately.

Physiological and anatomical responses to drought stress differ between two larch species and their hybrid

Key MessageHybrid saplings were more reactive to soil water deficit than Japanese and European larch. European larch had hydraulically safer wood and anisohydric behavior, Japanese and hybrid larch showed isohydric strategy.Deciduous larch species could be an alternative to evergreen conifers in reforestation, but little is known about drought sensitivity of their saplings. The effect of an experimental drought on hydraulics and quantitative wood anatomy was tested on saplings of European larch (EL, Larix decidua), Japanese larch (JL, Larix kaempferi) and their hybrid (HL). Across species, biomass, transpiration rate and relative water content were higher in controls than in drought stressed trees, but transpiration efficiency was lower. JL had the highest transpiration efficiency under drought, and EL the lowest, coinciding with slower growth of EL. Wood of EL formed before drought was hydraulically safer as shown by higher wall/lumen ratio and lower pit cavity area. EL neither had a significant increase in transpiration efficiency nor a reduction in transpiration rate under drought, suggesting that the stomata remained open under soil water deficit. HL saplings were the most reactive to water shortage, indicated by intra-annual density fluctuations and a decrease in relative water content of the sapwood. Significant reduction in transpiration by HL suggested a higher stomatal sensitivity, while the same leaf surface area was maintained and radial growth was still similar to its best parent, the JL. The latter showed a significantly lower leaf surface area under drought than controls. EL, with its hydraulically safer wood, followed an anisohydric behavior, while JL and HL revealed an isohydric strategy. Altogether, our results suggest species dependent acclimations to drought stress, whereby HL followed the strategy of JL rather than that of EL.

SabaTracheid 1.0: A Novel Program for Quantitative Analysis of Conifer Wood Anatomy - A Demonstration on African Juniper From the Blue Nile Basin.

Knowledge about past climates, especially at a seasonal time scale, is important as it allows informed decisions to be made to mitigate future climate change. However, globally, and especially in semi-arid Tropics, instrumental climatic data are scarce. A dendroclimatic approach may fill this gap, but tropical dendrochronological data are rare and do not yet provide fine resolution intra-annual information about past climates. Unlike in the Tropics, in the Mediterranean, temperate, alpine, and arctic regions, dendroanatomy and quantitative wood anatomy (QWA) are progressing fast attaining an intra-annual resolution, which allows a better understanding of seasonal climate dynamics and climate–growth relationships. The existing dendroanatomical and QWA methods aren’t suitable for tropical trees because they do not consider the high variation in tree ring width and the frequent occurrence of micro-rings containing only a few tracheids per radial file. The available tracheid analysis programs generally fail to provide multiple sectors for micro-rings and they are unable to compute most of the useful dendroanatomical parameters at fine temporal resolutions. Here, we present a program (SabaTracheid) that addresses the three main standard tasks that are necessary for QWA and dendroanatomy before running a climate analysis: (1) tracheidogram standardization, (2) sectoring, and (3) computing QWA and dendroanatomical variables. SabaTracheid is demonstrated on African Juniper (Juniperus procera Hochst. ex Endl), but it is potentially able to provide fine-resolution QWA and dendroanatomic data that could be used for dendroanatomical studies in all regions of the world. SabaTracheid is a freeware that quickly and accurately standardizes tracheidograms, divides tree rings into multiple regular sectors, computes useful dendroanatomic and QWA variables for the whole tree rings, early- and latewood portions, and each sector separately. This program is particularly adapted to deal with high inter-annual growth variations observed in tropical trees so that it assures the provision of complete sectoral QWA and dendroanatomical data for micro-rings as well. We demonstrate SabaTracheid using a dataset of 30 Juniperus procera tree rings from the Blue Nile basin, in Ethiopia. SabaTracheid’s ability to provide fine resolution QWA and dendroanatomic data will help the discipline develop in tropical as well as in the Mediterranean and temperate regions.

Canopy status modulates formation of wood rays in scots pine under hemiboreal conditions

Non-structural carbohydrates (NSC) reserves are crucial for trees to cope with weather extremes, thus to ensure their survival and ecological plasticity. The NSC reserves can depend on social status, suggesting uneven plasticity of trees at the stand level. In stemwood of Scots pine (Pinus sylvestris L.), which is a widespread and important species, NSC reserves are stored in parenchyma in wood rays (WR). The quantity of WRs is adjusted intra-annually, allowing retrospective analysis of factors affecting their formation. Accordingly, the differences in WR quantity in stemwood of dominant and intermediate (canopy trees with reduced and narrow crowns) maturing Scots pine were assessed by quantitative wood anatomy. Tangential cuts from the outermost 30 tree-rings were analysed. The relative ray area was intermediate, i.e. covering about 5% of the tangential cut, yet expressed high individuality among the trees. The size and amount of WR mainly differed between the earlywood and latewood; WRs in latewood were higher although narrower in comparison to earlywood, yet their total amount was higher in earlywood. Canopy status had only a slight effect, as quantity and height of WR tended to be higher for the intermediate trees, particularly in earlywood. The size and quantity of WR expressed inter-annual variation, which was mainly related to the meteorological conditions prior to the formation of the tree-ring (previous summer and autumn) indicating legacy effects of climatic factors on NSC and susceptibility of trees to cumulative effects of weather extremes. However, the climatic signals in the inter-annual variation of WR were weaker than observed before, likely due to location of the studied stand in the mid-part of the species range. Nevertheless, the observed differences in mean values and inter-annual variation of WR suggested a within-species diversity of carbon allocation patterns, supporting adaptability of the species.

Effects of Intra-Seasonal Drought on Kinetics of Tracheid Differentiation and Seasonal Growth Dynamics of Norway Spruce along an Elevational Gradient

Research Highlights: Our results provide novel perspectives on the effectiveness and collapse of compensatory mechanisms of tracheid development of Norway spruce during intra-seasonal drought and the environmental control of intra-annual density fluctuations. Background and Objectives: This study aimed to compare and integrate complementary methods for investigating intra-annual wood formation dynamics to gain a better understanding of the endogenous and environmental control of tree-ring development and the impact of anticipated climatic changes on forest growth and productivity. Materials and Methods: We performed an integrated analysis of xylogenesis observations, quantitative wood anatomy, and point-dendrometer measurements of Norway spruce (Picea abies (L.) Karst.) trees growing along an elevational gradient in South-western Germany during a growing season with an anomalous dry June followed by an extraordinary humid July. Results: Strong endogenous control of tree-ring formation was suggested at the highest elevation where the decreasing rates of tracheid enlargement and wall thickening during drought were effectively compensated by increased cell differentiation duration. A shift to environmental control of tree-ring formation during drought was indicated at the lowest elevation, where we detected absence of compensatory mechanisms, eventually stimulating the formation of an intra-annual density fluctuation. Transient drought stress in June also led to bimodal patterns and decreasing daily rates of stem radial displacement, radial xylem growth, and woody biomass production. Comparing xylogenesis data with dendrometer measurements showed ambivalent results and it appears that, with decreasing daily rates of radial xylem growth, the signal-to-noise ratio in dendrometer time series between growth and fluctuations of tree water status becomes increasingly detrimental. Conclusions: Our study provides new perspectives into the complex interplay between rates and durations of tracheid development during dry-wet cycles, and, thereby, contributes to an improved and mechanistic understanding of the environmental control of wood formation processes, leading to the formation of intra-annual density fluctuations in tree-rings of Norway spruce.

Intra‐Annual Climate Anomalies in Northwestern North America Following the 1783–1784 CE Laki Eruption

AbstractThe 1783–1784 CE Laki eruption in Iceland was one of the largest, in terms of the mass of SO2 emitted, high‐latitude eruptions in the last millennium, but the seasonal and regional climate response was heterogeneous in space and time. Although the eruption did not begin until early June, tree‐ring maximum latewood density (MXD) reconstructions from Alaska suggest that the entire 1783 summer was extraordinarily cold. We use high‐resolution quantitative wood anatomy, climate model simulations, and proxy systems modeling to resolve the intra‐annual climate effects of the Laki eruption on temperatures over northwestern North America. We measured wood anatomical characteristics of white spruce (Picea glauca) trees from two northern Alaska sites. Earlywood cell characteristics of the 1783 ring are normal, while latewood cell wall thickness is significantly and anomalously reduced compared to non‐eruption years. Combined with complementary evidence from climate model experiments and proxy systems modeling, these features indicate an abrupt and premature cessation of cell wall thickening due to a rapid temperature decrease toward the end of the growing season. Reconstructions using conventional annual resolution MXD likely over‐estimate total growing season cooling in this year, while ring width fails to capture this abrupt late‐summer volcanic signal. Our study has implications not only for the interpretation of the climatic impacts of the Laki eruption in North America, but more broadly demonstrates the importance of timing and internal variability when comparing proxy temperature reconstructions and climate model simulations. It further demonstrates the value of developing cellular‐scale tree‐ring proxy measurements for paleoclimatology.