Developmental and environmental factors driving xylem anatomy and micro-density

Author(s):  
Annie Deslauriers ◽  
Valentinà Buttò ◽  
Fabrizio Cartenì ◽  
Sergio Rossi ◽  
Hubert Morin

<p>The current research on the dynamics of tree ring formation in conifers has provided new insights into how rate and duration xylem-cell production and development control the size of the xylem conduits leading to the formation of earlywood and latewood. So far, the physiology behind wood formation processes and the associated kinetics has rarely been considered, leading to the impossibility to grasp the drivers of wood density changes along the tree-rings. Despite the importance of wood density for carbon sequestration and tree hydraulics, little is known about the factors controlling variations in wood density across the tree ring, i.e. micro-density, at the intra-annual scale. We first developed a process-based mechanistic model that simulates the development of conifer tracheids from a simple sugar signal that we discuss together with the main kinetics and environmental variables leading to the formation of micro-density in black spruce, the main conifers species in the boreal forest of Canada. At the beginning of the growing season, low sugar availability in the cambium results in slow wall deposition that allows for a lengthier enlargement time thus producing large cells with thin walls (i.e. earlywood). In late summer and early autumn, high sugar availability produces narrower cells with thick cell walls (i.e. latewood). Wood formation dynamics had an indirect effect on micro-density. Micro-density increased under longer periods of cell wall deposition and shorter durations of enlargement. Cell diameter indirectly affected micro-density via cell wall thickness, which was the most important parameter affecting micro-density. Cell traits experienced the joint action of enlargement and secondary wall deposition in shaping the intra-annual patterns of tree rings. Our results point to the predictive power of a simple sugar signal. During the growing season, the amount of carbon allocated to wood formation largely influences the duration of cell differentiation, thus modulating cell diameter, cell wall thickness and by result tree-ring micro-density.</p><p> </p>

Holzforschung ◽  
2013 ◽  
Vol 67 (2) ◽  
pp. 217-225 ◽  
Author(s):  
Jinmei Xu ◽  
Jianxiong Lu ◽  
Fucheng Bao ◽  
Robert Evans ◽  
Geoffrey M. Downes

Abstract Dimensions of dated tree rings are an important tool of dendroclimatology. However, the relationships between climatic variables and cell diameter and cell wall thickness are not yet clearly elaborated. In the present article, year-to-year cell characteristics, ring width, and wood density of Picea crassifolia trees growing in northwestern China have been measured with high resolution by means of the instrument SilviScan-3. The response function analysis showed that climate explained 51% of the variation of cell radial diameter chronology, 48% of wood density, 40% of cell wall thickness, and 37% of ring width. Cell wall thickness and wood density responded significantly and positively to temperature, and the response to precipitation was negative, while the opposite was true for cell radial diameter and ring width. Cell wall thickness and wood density were pronounced (statistically significant) to temperature in September and precipitation in May and August. Cell radial diameter responded significantly to temperature in June and July, and precipitation, in August. For ring width, the temperature in July was important. Accordingly, cell characteristics are sensitive to climate, and the findings could be useful in the field of dendroclimatology.


2021 ◽  
Author(s):  
Milagros Rodriguez-Caton ◽  
Laia Andreu-Hayles ◽  
Mariano S Morales ◽  
Valérie Daux ◽  
Duncan A Christie ◽  
...  

Abstract Tree growth is generally considered to be temperature-limited at upper elevation treelines. Yet, climate factors controlling tree growth at semiarid treelines are poorly understood. We explored the influence of climate on stem growth and stable isotopes for Polyepis tarapacana, the world’s highest elevation tree-species found only in the South American Altiplano. We developed tree-ring width index (RWI), oxygen (δ18O) and carbon (δ13C) chronologies for the last 60 years at four P. tarapacana stands located above 4,400 meters in elevation, along a 500-km latitude-aridity gradient. Total annual precipitation decreased from 300 to 200 mm from the northern to the southern sites. We used RWI as a proxy of wood formation (carbon sink) and isotopic tree-ring signatures as proxies of leaf-level gas exchange processes (carbon source). We found distinct climatic conditions regulating carbon-sink processes along the gradient. Current-growing season temperature regulated RWI at wetter-northern sites, while prior-growing season precipitation determined RWI at arid-southern sites. This suggests that the relative importance of temperature to precipitation in regulating tree growth is driven by site-water availability. In contrast, warm and dry growing-seasons resulted in enriched tree-ring δ13C and δ18O at all study sites, suggesting that similar climate conditions control carbon-source processes. Site-level δ13C and δ18O chronologies were significantly and positively related at all sites, with the strongest relationships among the southern-drier stands. This indicates an overall regulation of intercellular carbon dioxide via stomatal conductance for the entire P. tarapacana network, with greater stomatal control when aridity increases. The manuscript also highlights a coupling and decoupling of physiological processes at leaf level versus wood formation depending on their respectively uniform and distinct sensitivity to climate. This study contributes to better understand and predict the response of high-elevation Polylepis woodlands to rapid climate changes and projected drying in the Altiplano.


2015 ◽  
Vol 40 (2) ◽  
pp. 315-340 ◽  
Author(s):  
B. A. Molski

The corewood of pine ds very prone to compression wood formation, this changing the whole pattern of the tree ring structure and the siz.es of early and late wood. Compression wood always increases the formation of late wood at the expense of early wood. Tree rings with compression wood are generally wider than those without it, but there occur also tree rings wihout compression wood wider than those in which it is present, formed in the same year and in the same tree.


Forests ◽  
2020 ◽  
Vol 11 (2) ◽  
pp. 146 ◽  
Author(s):  
Franco Biondi

The contribution of tree-ring analysis to other fields of scientific inquiry with overlapping interests, such as forestry and plant population biology, is often hampered by the different parameters and methods that are used for measuring growth. Here I present relatively simple graphical, numerical, and mathematical considerations aimed at bridging these fields, highlighting the value of crossdating. Lack of temporal control prevents accurate identification of factors that drive wood formation, thus crossdating becomes crucial for any type of tree growth study at inter-annual and longer time scales. In particular, exactly dated tree rings, and their measurements, are crucial contributors to the testing and betterment of allometric relationships.


2019 ◽  
Vol 123 (7) ◽  
pp. 1257-1265 ◽  
Author(s):  
Valentina Buttò ◽  
Sergio Rossi ◽  
Annie Deslauriers ◽  
Hubert Morin

Abstract Background and Aims Secondary growth is a process related to the formation of new cells that increase in size and wall thickness during xylogenesis. Temporal dynamics of wood formation influence cell traits, in turn affecting cell patterns across the tree ring. We verified the hypothesis that cell diameter and cell wall thickness are positively correlated with the duration of their differentiation phases. Methods Histological sections were produced by microcores to assess the periods of cell differentiation in black spruce [Picea mariana (Mill.) B.S.P.]. Samples were collected weekly between 2002 and 2016 from a total of 50 trees in five sites along a latitudinal gradient in Quebec (Canada). The intra-annual temporal dynamics of cell differentiation were estimated at a daily scale, and the relationships between cell traits and duration of differentiation were fitted using a modified von Bertalanffy growth equation. Key Results At all sites, larger cell diameters and cell wall thicknesses were observed in cells that experienced a longer period of differentiation. The relationship was a non-linear, decreasing trend that occasionally resulted in a clear asymptote. Overall, secondary wall deposition lasted longer than cell enlargement. Earlywood cells underwent an enlargement phase that lasted for 12 d on average, while secondary wall thickness lasted 15 d. Enlargement in latewood cells averaged 7 d and secondary wall deposition occurred over an average of 27 d. Conclusions Cell size across the tree ring is closely connected to the temporal dynamics of cell formation. Similar relationships were observed among the five study sites, indicating shared xylem formation dynamics across the entire latitudinal distribution of the species.The duration of cell differentiation is a key factor involved in cell growth and wall thickening of xylem, thereby determining the spatial variation of cell traits across the tree ring.


Atmosphere ◽  
2020 ◽  
Vol 11 (3) ◽  
pp. 247
Author(s):  
Zhipeng Dong ◽  
Dai Chen ◽  
Jianhua Du ◽  
Guang Yang ◽  
Maowei Bai ◽  
...  

Humid subtropical China is an “oasis” relative to other dry subtropics of the world due to the prevailing of the East Asian summer monsoon (EASM). Although many long climate sensitive tree-rings have been published to understand the historical climate change over various regions in China, long tree-ring chronologies in humid subtropical China are rare due to the difficulty to find old growth trees. This study established a tree-ring chronology spanning from 1776 to 2016 from Cryptomeria fortunei Hooibrenk ex Otto et Dietr in Liancheng area of humid subtropical China, which is also currently the longest chronology in Fujian province. Similar to the climate-growth relationships in neighboring regions, our tree-ring chronology is limited by cold temperature in winter and spring and drought in summer. In addition, a drought stress before the growing season also played a role in limiting the growth of our tree rings. Our climate sensitive tree rings showed different correlations with the Pacific Decadal Oscillation (PDO) in different periods, possibly via modulation of the EASM.


Forests ◽  
2020 ◽  
Vol 11 (12) ◽  
pp. 1294
Author(s):  
Liliana V. Belokopytova ◽  
Patrick Fonti ◽  
Elena A. Babushkina ◽  
Dina F. Zhirnova ◽  
Eugene A. Vaganov

Research Highlights: This study emphasized the importance of multi-parameter analyses along ecological gradients for a more holistic understanding of the complex mechanism of tree-ring formation. Background and Objectives: The analysis of climatic signals from cell anatomical features measured along series of tree-rings provides mechanistic details on how environmental drivers rule tree-ring formation. However, the processes of cell development might not be independent, limiting the interpretation of the cell-based climatic signal. In this study, we investigated the variability, intercorrelations and climatic drivers of wood anatomical parameters, resulting from consequent cell developmental processes. Materials and Methods: The study was performed on thin cross-sections from wood cores sampled at ~1.3 m stem height from mature trees of Pinus sylvestris L. growing at five sampling sites along an ecological gradient from cold and wet to hot and dry within continental Southern Siberia. Tracheid number per radial file, their diameters and wall thicknesses were measured along the radial direction from microphotographs for five trees per site. These parameters were then averaged at each site for earlywood and latewood over the last 50 tree rings to build site chronologies. Their correlations among themselves and with 21-day moving climatic series were calculated. Results: Our findings showed that wood formation was not simply the result of environmentally driven independent subprocesses of cell division, enlargement and wall deposition. These processes appear to be interconnected within each zone of the ring, as well as between earlywood and latewood. However, earlywood parameters tend to have more distinctive climatic responses and lower intercorrelations. On the other hand, there are clear indications that the mechanisms of cell division and enlargement share similar climatic drivers and are more sensitive to water limitation than the process of wall deposition. Conclusions: Indications were provided that (i) earlywood formation left a legacy on latewood formation, (ii) cell division and enlargement shared more similar drivers between each other than with wall deposition, and (iii) the mechanism of cell division and enlargement along the gradient switch from water to heat limitations at different thresholds than wall deposition.


1971 ◽  
Vol 1 (4) ◽  
pp. 419-449 ◽  
Author(s):  
Harold C. Fritts

Dendrochronology is the science of dating annual growth layers (rings) in woody plants. Two related subdisciplines are dendroclimatology and dendroecology. The former uses the information in dated rings to study problems of present and past climates, while the latter deals with changes in the local environment rather than regional climate.Successful applications of dendroclimatology and dendroecology depend upon careful stratification. Ring-width samples are selected from trees on limiting sites, where widths of growth layers vary greatly from one year to the next (sensitivity) and autocorrelation of the widths is not high. Rings also must be cross-dated and sufficiently replicated to provide precise dating. This selection and dating assures that the climatic information common to all trees, which is analogous to the “signal”, is large and properly placed in time. The random error or nonclimatic variations in growth, among trees, is analogous to “noise” and is reduced when ring-width indices are averaged for many trees.Some basic facts about the growth are presented along with a discussion of important physiological processes operating throughout the roots, stems, and leaves. Certain gradients associated with tree height, cambial age, and physiological activity control the size of the growth layers as they vary throughout the tree. These biological gradients interact with environmental variables and complicate the task of modeling the relationships linking growth with environment.Biological models are described for the relationships between variations in ring widths from conifers on arid sites, and variations in temperature and precpitation. These climatic factors may influence the tree at any time in the year. Conditions preceding the growing season sometimes have a greater influence on ring width than conditions during the growing season, and the relative effects of these factors on growth vary with latitude, altitude, and differences in factors of the site. The effects of some climatic factors on growth are negligible during certain times of the year, but important at other times. Climatic factors are sometimes directly related to growth and at other times are inversely related to growth. Statistical methods are described for ascertaining these differences in the climatic response of trees from different sites.A practical example is given of a tree-ring study and the mechanics are described for stratification and selection of tree-ring materials, for laboratory preparation, for cross-dating, and for computer processing. Several methods for calibration of the ring-width data with climatic variation are described. The most recent is multivariate analysis, which allows simultaneous calibration of a variety of tree-ring data representing different sites with a number of variables of climate.Several examples of applications of tree-ring analysis to problems of environment and climate are described. One is a specification from tree rings of anomalies in atmosphere circulation for a portion of the Northern Hemisphere since 1700 A.D. Another example treats and specifies past conditions in terms of conditional probabilities. Other methods of comparing present climate with past climate are described along with new developments in reconstructing past hydrologic conditions from tree rings.Tree-ring studies will be applied in the future to problems of temperate and mesic environments, and to problems of physiological, genetic, and anatomical variations within and among trees. New developments in the use of X-ray techniques will facilitate the measurement and study of cell size and cell density. Tree rings are an important source of information on productivity and dry-matter accumulation at various sites. Some tree-ring studies will deal with environmental pollution. Statistical developments will improve estimation of certain past anomalies in weather factors and the reconstructtion of atmosphere circulation associated with climate variability and change. Such information should improve chances for measuring and assessing the possibility of inadvertent modification of climate by man.


2020 ◽  
Author(s):  
Gonzalo Peres-De-Lis ◽  
Cyrille Rathgeber ◽  
Stéphane Ponton

<p>Despite a long-standing interest in retrieving intra-annual environmental information from tree-ring features, none of the approaches developed so far for accurately dating intra-ring sector has been validated on observations. Here, we investigated space-for-time association across regular intra-ring sectors for which we estimated the timing of formation. For this purpose, a unique dataset containing quantitative wood anatomy measurements and kinetics of tracheid differentiation was compiled for 45 trees grown in North-East France (three years of wood formation monitoring, for five trees, from three different conifer species). Tracheid dimensions were measured directly on the best anatomical sections at the end of the growing season, while the kinetics of xylem cell differentiation were provided at tree-level by an empirical model of wood formation dynamics. Our results confirmed that the time taken to form sectors of the same width increased from earlywood (composed of wide thin-walled tracheids) to latewood (composed of narrow thick-walled tracheids). This mainly reflected the increase of the duration of cell wall deposition through the growing season, and, to a lesser extent, the augmentation of the number of tracheids per sectors. However, our results also show that regular intra-ring sectors, which were well separated in space, overlapped in time. The overlapping culminated during the summer period, reaching 40 % for 10 sectors. It could be reduced to approx. 30 % by increasing the number of sectors (from 10 to 25, for example), but it cannot be removed. Therefore, successive intra-ring sectors could not be attributed to a succession of separated time intervals by simply using their relative position along the ring. However, the formation of sectors of equivalent ranks were noticeably synchronous between the different trees and years, reaching 80 % of synchronicity for the process of wall thickening. This suggest that data from regular intra-ring sectors could be reliably used to build mean chronologies expressing the common signal of tree populations. Our results show the limits that the xylogenesis process itself imposes on the dating of intra-ring features. They also argue for an in-depth understanding of the association between cell differentiation processes (enlargement, wall thickening and lignification) and wood characteristics (density, anatomy, stable isotope composition).</p>


IAWA Journal ◽  
2019 ◽  
Vol 40 (2) ◽  
pp. 155-182 ◽  
Author(s):  
Veronica De Micco ◽  
Marco Carrer ◽  
Cyrille B.K. Rathgeber ◽  
J. Julio Camarero ◽  
Jordi Voltas ◽  
...  

ABSTRACTIt is noteworthy that the largest part of global vegetation biomass depends on a thin layer of cells: the vascular cambium. Understanding the wood formation processes and relationships with environmental factors is a crucial and timely research question requiring interdisciplinary efforts, also to upscale the information gained and to evaluate implications for tree growth and forest productivity.We provide an overview of wood formation processes up to tree-ring development, bearing in mind that the combined action of intrinsic factors and environmental drivers determines the anatomical traits of a tree ring formed at a specific time and position within the tree’s architecture. After briefly reviewing intrinsic factors, we focus attention on environmental drivers highlighting how a correct interpretation of environmental signals in tree rings must be grounded in a deep knowledge of xylogenesis and consequent wood anatomical traits. We provide guidelines on novel methods and approaches recently developed to study xylem formation. We refer to existing literature on established techniques for retrospective analyses in tree-ring series of anatomical and isotopic traits, to assess long-term ecophysiological responses to environmental variations, also giving advice on possible bias because of interand within-tree variability.Finally, we highlight that, once the temporal axis of intra-annual tree-ring variability of xylem traits is established by xylogenesis analysis, a multidisciplinary approach linking classical dendro-ecology, wood functional traits (dendro-anatomy) and eco-physiology (here focusing on dendro-isotopes) allows a better interpretation of past environmental events hidden in tree rings, and more reliable forecasts of wood growth in response to climate change.


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