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.
ABSTRACT Anatomical and subcellular characteristics of juvenile and adult wood of seven species (Manilkara zapota (L.) P. Royen, Platymiscium yucatanum Standl., Lonchocarpus castilloi Standl., Roseodendron donnell-smithii (Rose) Miranda, Terminalia buceras (L.) Wright, Tabebuia rosea (Bertol.) DC., Lysiloma latisiliquum (L.) Benth. from Mexico, including a histometric evaluation, were investigated by light microscopy with a digitized image analysis system and by X-ray diffractometry using the SilviScan® system. The topochemical distribution of lignin and phenolic deposits in the tissue was studied by means of cellular UV-microspectrophotometry (UMSP). Extractive contents (acetone/water and water) were determined gravimetrically. The results of the structural and topochemical analyses were compared with the interrelations of certain anatomical and subcellular structures as well as the topochemical composition with regard to the physical and mechanical properties of the timbers investigated. The objective was to provide a detailed cellular and subcellular description of the heartwood of seven lesser-known timbers from Central America. All examined tree species show significant differences between juvenile and adult heartwood. For individual species, however, the differences vary greatly and have to be individually assessed in addition to general trends observed for all studied species. It has been shown that vessel size, fibre length, size of fibre lumina, and height and width of rays, as well as the content of extractives and topochemical composition of the cell walls, are suitable indicators for the differentiation of the two heartwood types. The results also contribute to a better understanding of the wood properties of the investigated timbers in relation to their utilization and added value increase.
ABSTRACTRelationships between hydraulic vulnerability expressed asP50(the air pressure causing 50% loss of hydraulic conductivity) and within-ring differences in wood density (WD) and anatomical features were investigated with the aim to find efficient proxies forP50relating to functional aspects. WD and tracheid dimensions were measured with SilviScan on Norway spruce (Picea abies(L.) Karst.) trunk wood.P50was strongly related to mean WD (r= -0.64) and conduit wall reinforcement ((t/b)2), the square of the ratio between the tracheid double wall thickness (t) and the lumen width (b), where use of tangential lumen width ((t/bt)2) gave better results (r= -0.54) than radial lumen width (r= -0.31). The correlations ofP50with earlywood (EW), transition wood (TW) and latewood (LW) traits were lower than with the specimen averages, both for WD (r= -0.60 for WDEW,r= -0.56 for WDTW,r= -0.23 for WDLW) and all anatomical traits. The loss of hydraulic conductivity was addressed as a dynamic process and was simulated by defining consecutive phases of 5% theoretical conductivity loss. WD and tracheid traits were calculated and correlated withP50values of each specimen. Tightest correlations were found for (t/bt)2, at relative cumulated theoretical conductivities until 45 to 50% (r= -0.75).We conclude that WD is one of the best available proxies forP50, but does not necessarily reflect the mechanism behind resistance to cavitation. The new trait, based on estimation of conductivity loss as a dynamic process, provided even stronger correlations.
To maximize utilization of our forest resources, detailed knowledge of wood property variation and the impacts this has on end-product performance is required at multiple scales (within and among trees, regionally). As many wood properties are difficult and time-consuming to measure our knowledge regarding their variation is often inadequate as is our understanding of their responses to genetic and silvicultural manipulation. The emergence of many non-destructive evaluation (NDE) methodologies offers the potential to greatly enhance our understanding of the forest resource; however, it is critical to recognize that any technique has its limitations and it is important to select the appropriate technique for a given application. In this review, we will discuss the following technologies for assessing wood properties both in the field: acoustics, Pilodyn, Resistograph and Rigidimeter and the lab: computer tomography (CT) scanning, DiscBot, near infrared (NIR) spectroscopy, radial sample acoustics and SilviScan. We will discuss these techniques, explore their utilization, and list applications that best suit each methodology. As an end goal, NDE technologies will help researchers worldwide characterize wood properties, develop accurate models for prediction, and utilize field equipment that can validate the predictions. The continued advancement of NDE technologies will also allow researchers to better understand the impact on wood properties on product performance.
Very few annually resolved millennial-length temperature reconstructions exist for the Southern Hemisphere. Here we present four 979-year reconstructions for southeastern Australia for the austral summer months of December–February. Two of the reconstructions are based on the Australian Water Availability Project dataset and two on the Berkeley Earth Surface Temperature dataset. For each climate data set, one reconstruction is based solely on Lagarostrobos franklinii (restricted reconstructions) while the other is based on multiple Tasmanian conifer species (unrestricted reconstructions). Each reconstruction calibrates ~50−60% of the variance in the temperature datasets depending on the number of tree-ring records available for the reconstruction. We found little difference in the temporal variability of the reconstructions, although extremes are amplified in the restricted reconstructions relative to the unrestricted reconstructions. The reconstructions highlight the occurrence of numerous individual years, especially in the 15th−17th Centuries, for which temperatures were comparable with those of the late 20th Century. The 1950−1999 period, however, stands out as the warmest 50-year period on average for the past 979 years, with a sustained shift away from relatively low mean temperatures, the length of which is unique in the 979-year record. The reconstructions are strongly and positively related to temperatures across the southeast of the Australian continent, negatively related to temperatures in the north and northeast of the continent, and uncorrelated with temperatures in the west. The lack of a strong relationship with temperatures across the continent highlights the necessity of a sub-regional focus for Australasian temperature reconstructions.
Seasonal variability is a significant source of uncertainty in projected changes to precipitation across southeastern Australia (SEA). While existing instrumental records provide seasonal data for recent decades, most proxy records (e.g., tree rings, corals, speleothems) offer only annual reconstructions of hydroclimate. We present the first cool‐season (July–August) reconstruction of dam inflow (Lake Burbury) for western Tasmania in SEA based on tree‐ring width ( Athrotaxis selaginoides ) and mean latewood cell wall thickness ( Phyllocladus aspleniifolius ) chronologies. The reconstruction, produced using principal component regression, verifies back to 1731 and is moderately skillful, explaining around 23% of the variance. According to the reconstruction, relatively low inflow periods occurred around 1860, the early 1900s and 1970, while relatively high inflows occurred in the 1770s and 1810s. Highest reconstructed inflows occurred in 1816, and lowest in 1909. Comparison with available documentary and instrumental records indicates that the reconstruction better captures high rather than low flow events. There is virtually no correlation between our reconstruction and another for December–January inflow for the same catchment, a result consistent with the relationship between seasonal instrumental data. This suggests that conditions in one season have not generally reflected conditions in the other season over the instrumental record, or for the past 277 years. This illustrates the value of obtaining reconstructions of regional hydroclimatic variability for multiple individual seasons in regions where dry and wet seasons are not strongly defined. The results also indicate that the hydroclimate of the southeastern Australian region cannot be adequately represented by a single regional reconstruction.
More than one-third of Germany s Douglas-fir resources, stock in age-classes from twenty-one to fifty-nine years. As such timber increasingly enters markets, detailed knowledge of the anatomy and properties of its wood is of importance to forest managers and wood processors. Anatomical and mechanical wood analyses in this study were carried out on twenty trees from four scientifically managed plantations in Southern Germany. The age of the trees selected was forty-two years whereby varying growth conditions were considered. Juvenile and adult woods were identified by segmented linear regression of radial profiles of anatomical characteristics, such as latewood percentage, tracheid wall thickness microfibril angle and density. Additionally, the width of earlywood, latewood and growth rings as well as bending modulus of elasticity were determined. Variance was dependent on the trait used for differentiation, j uvenile wood comprised of an eleven to thirty-one growth rings resp. radial amounts of fifteen to sixty-five percent. When compared to adult wood, juvenile wood showed corresponding features of approximately thirty percent wider growth rings, th irty four percent lower latewood percentage, fourteen percent thinner tracheid walls, and eighty percent larger microfibril angles, eleven percent lighter wood and fifty -seven percent lower bending modulus of elasticity. As the assortment features fast grown trees, adult heartwood characteristics were slightly inferior to the characteristics of European Douglas-fir.
Tree-ring based climate reconstructions have been critical for understanding past variability and recent trends in climate worldwide, but they are scarce in Australia. This is, particularly the case for temperature: only one tree ring width based temperature reconstruction - based on Huon Pine trees from Mt Read, Tasmania - exists for Australia. Here, we investigate whether additional tree-ring parameters derived from Athrotaxis cupressoides trees growing in the same region have potential to provide robust proxy records of past temperature variability. We measured wood properties, including tree-ring width (TRW), mean density, mean cell wall thickness (CWT), and tracheid radial diameter (TRD) of annual growth rings in Athrotaxis cupressoides, a long-lived, high-elevation conifer in central Tasmania, Australia. Mean density and CWT were strongly and negatively correlated with summer temperatures. In contrast, the summer temperature signal in TRW was weakly positive. The strongest climate signal in any of the tree-ring parameters was maximum temperature in January (mid-summer; JanT(max)) and we chose this as the target climate variable for reconstruction. The model that explained most of the variance in JanTmax was based on TRW and mean density as predictors. TRW and mean density provided complementary proxies with mean density showing greater high-frequency (inter-annual to multi-year) variability and TRW showing more low-frequency (decadal to centennial-scale) variability. The final reconstruction model is robust, explaining 55% of the variance in JanT(max), and was used to reconstruct JanT(max) for the last five centuries (1530-2010 C.E.). The reconstruction suggests that the most recent 60 years have been warmer than average in the context of the last ca. 500 years. This unusually warm period is likely linked to a coincident increase in the intensity of the subtropical ridge and dominance of the positive phase of the Southern Annular Mode in summer, which weaken the influence of the band of prevailing westerly winds and storms on Tasmanian climate. Our findings indicate that wood properties, such as mean density, are likely to provide significant contributions toward the development of robust climate reconstructions in the Southern Hemisphere and thus toward an improved understanding of past climate in Australasia. (C) 2016 Elsevier B.V. All rights reserved.
There is evidence that recently occurring top dieback of Norway spruce (Piceaabies(L.) Karst.) treesin southern Norway is associated with drought stress. We comparedfunctional wood traits of20 healthy looking trees and 20 trees with visual signs of top dieback. SilviScan technology was applied to measure cell dimensions (lumen and cell wall thickness) in a selected set of trunk wood specimens where vulnerability to cavitation (P50) datawere available. The conduit wall reinforcement ((t/b)²) was a good proxy for P50. Cell dimensions were measured on wood cores of all 40 trees; theoretical vulnerability of single annual rings could bethus estimated. Declining trees tended to have lower (t/b)² before and during a period of water deficit (difference between precipitation and potential evapotranspiration)that lasted from 2004 to 2006. The results are discussed with respect to genetic predisposition.
Top dieback in 40-60 years old forest stands of Norway spruce [Picea abies (L.) Karst.] in southern Norway is supposed to be associated with climatic extremes. Our intention was to learn more about the processes related to top dieback and in particular about the plasticity of possible predisposing factors. We aimed at (i) developing proxies for P50 based on anatomical data assessed by SilviScan technology and (ii) testing these proxies for their plasticity regarding climate, in order to (iii) analyze annual variations of hydraulic proxies of healthy looking trees and trees with top dieback upon their impact on tree survival. At two sites we selected 10 tree pairs, i.e., one healthy looking tree and one tree with visual signs of dieback such as dry tops, needle shortening and needle yellowing (n = 40 trees). Vulnerability to cavitation (P50) of the main trunk was assessed in a selected sample set (n = 19) and we thereafter applied SilviScan technology to measure cell dimensions (lumen (b) and cell wall thickness (t)) in these specimen and in all 40 trees in tree rings formed between 1990 and 2010. In a first analysis step, we searched for anatomical proxies for P50. The set of potential proxies included hydraulic lumen diameters and wall reinforcement parameters based on mean, radial, and tangential tracheid diameters. The conduit wall reinforcement based on tangential hydraulic lumen diameters ((t/bht)2) was the best estimate for P50. It was thus possible to relate climatic extremes to the potential vulnerability of single annual rings. Trees with top dieback had significantly lower (t/bht)2 and wider tangential (hydraulic) lumen diameters some years before a period of water deficit (2005-2006). Radial (hydraulic) lumen diameters showed however no significant differences between both tree groups. (t/bht)2 was influenced by annual climate variability; strongest correlations were found with precipitation in September of the previous growing season: high precipitation in previous September resulted in more vulnerable annual rings in the next season. The results are discussed with respect to an "opportunistic behavior" and genetic predisposition to drought sensitivity.
Projected decreases and changes in the seasonal distribution of precipitation will have profound impacts on southeastern Australia, including its ability to generate renewable hydroelectricity. Recent decreases in precipitation over the region may be significant in the context of instrumental records, but the question of whether these decreases are within long‐term natural variability remains. To help address this issue, we present December–January streamflow and dam inflow reconstructions for southeastern Australia. These reconstructions for the Tasmanian west coast are based solely on local tree ring chronologies and span up to 1600 years. Nonparametric estimates, however, indicate good model skill for the last 458 years (streamflow) and 478 years (dam inflow). The reconstructions indicate that twentieth century conditions were well within the range of historical variability, and were in fact relatively wet. The period from approximately 1600 to 1750 CE was one of the enhanced variability and a high proportion of low and high flow events occurred in the seventeenth century. There are significant relationships between streamflow and inflow reconstructions and large‐scale ocean‐atmosphere processes such as ENSO and the Southern Annular Mode. Critically, our two reconstructions rely heavily on new tree ring chronologies based on properties such as tracheid radial diameter, cell wall thickness, and density, underscoring the importance of these different types of chronologies in reconstructions.
Little attention was given to the climatic signals in wood properties, such as microfibril angle (MFA) and tracheid radial diameter (TRD). In this article, year-to-year variation in MFA and TRD was measured by SilviScan-3 on dated Picea crassifolia trees growing at three altitudes in the northeastern Tibetan plateau. Climatic signals registered in MFA and TRD were analyzed using dendroclimatology methods. The annual variation of MFA and TRD was strongly linked to high-frequency climatic signals. Both MFA and TRD were negatively correlated with temperature and positively correlated with precipitation. The temperature had a similar influence on MFA and TRD at three different altitudes, while the influence of precipitation decreased with the increasing altitudes. MFA was negatively correlated with TRD, and this MFA–TRD internal relationship (R t) varied with calendar year. Temperature and precipitation had a strong influence on R t. Temperature was positively correlated with R t, and precipitation was negatively correlated with R t. The influence of temperature was stronger than that of precipitation. The influence of temperature increased with the increase in altitudes, while the influence of precipitation decreased with the increasing altitudes. Results of this study revealed that the trees could change their internal characteristics to adapt to the changing climate.
Little attention has been given to climatic signals in wood properties. In this study, ring width (RW), annual average microfibril angle (MFA), annual average tracheid radial diameter (TRD), and annual average density (DEN), as the annual and intra-annual wood property variables, were measured at high resolution by SilviScan-3 on dated Picea crassifolia trees. Dendroclimatological methods were used to analyze climatic signals registered in wood property variables. RW, MFA, and TRD negatively correlated with temperature and positively correlated with precipitation in the growing season, whereas the reverse was true for DEN. Climatic signals recorded in the earlywood were similar to those measured for the full width of the annual rings. Climatic signals recorded in latewood were very weak except for latewood MFA. This study showed that wood property variables could be extensive resources for learning more about the influences of climate on tree growth and how trees adapt to ongoing climate change.
In order to investigate the climate response of wood cell characteristics of Picea crossifolia at three elevations in the Qilian Mountains,northwestern China,we measured radial diameter,number and wall thickness of tracheids of P. crossifolia using Silviscan-3. Residual chronologies of radial diameter,number and wall thickness of tracheids were established by dendrochronological methods based on variation of cell characteristics with elevation and calendar year. Relationships between cell characteristics and monthly mean temperature and monthly total precipitation were analyzed. The results indicated that radial diameter of tracheids increased but the number of tracheids decreased with the rise of elevation; there was no obvious change in wall thickness at three elevations. Radial diameter of tracheids was negatively correlated with temperature,but positively associated with precipitation. The number of tracheids was positively correlated with temperature,but negatively with precipitation. Temperature in June and July,and precipitation in May and June had a significant influence on radial diameter and number of tracheids. Tracheid wall thickness was positively correlated with temperature,but negatively with precipitation. Temperature had a stronger effect on tracheid wall thickness than precipitation. Effect of temperature on cell characteristics was consistent at three elevations,while the impact of precipitationdecreased with the rise of elevation. Our findings suggest that precipitation,instead of temperature,is the main factor restricting the distribution of P. crossifolia along elevation gradient.
The formation of reaction wood is an adaptive feature of trees in response to various mechanical forces. In gymnosperms, reaction wood consists of compression wood (CW) and opposite wood (OW) that are formed on the underside and upperside of bent trunks and branches. Although reaction wood formed in bent trunks has been extensively investigated, relatively little has been reported from conifer branches. In this study SilviScan® technology was used to characterize radiata pine branches at high resolution. Compared to OW formed in the branches, CW showed greater growth, darker colour, thicker tracheid walls, higher coarseness, larger microfibril angle (MFA), higher wood density, lower extensional stiffness and smaller internal specific surface area. However, tracheids of CW were similar to those of OW in their radial and tangential diameters. These results indicated that gravity influenced tracheid cell division and secondary wall formation but had limited impact on primary wall expansion. Furthermore, seasonal patterns of CW formation were not observed in the branches from cambial age 4 while earlywood and latewood were clearly separated in all rings of OW. The marked change of MFA during reaction wood formation suggested that branches could be ideal materials for further study of cellulose microfibril orientation.
Ring width represents the total width of cells in a growing season. Relatively few studies have identified which of the parameters, cell number, size, or wall thickness is the greater contributor to variations in ring width. It is difficult to clearly explain the relationship between the climate in which a tree is located and the ring growth. In this study, the ring width and various tracheid characteristics (radial lumen diameter, double wall thickness, and number) in 298 rings of Picea crassifolia wood were investigated. Tree rings were quantitatively categorized based on their width into large, medium, and narrow classes. The tracheid number and lumen diameter were strongly correlated with annual ring width. The tracheid number had the strongest effect on the ring width. The effect of the tracheid number was three times larger than that of the lumen diameter. More earlywood cells were formed in larger rings, while larger earlywood cells were produced in years when narrow rings were formed. Wall thickness had no appreciable effect on ring width. The results of this study help to understand the relationship between the climate and the ring growth from the tree physiology perspective, when ring width is used as a climate proxy.
A radial core from a Norway spruce (Picea abies (L.) Karst.) estimated to be about 107 years old was cut from a board and was analyzed for density and microfibril angle (MFA). Furthermore, cell geometry, wall thickness and lignin distribution were analyzed on three selected growth rings in detail. Intra-ring differences in the density profiles are also true for cell wall thicknesses as well as radial and tangential lumen diameters. A higher MFA was found for earlywood with a slow decrease toward the latewood region. The lignin was found to remain rather constant throughout the growth rings, which suggests a constant chemical composition of the cell wall material within the growth ring. From the recorded datasets on a cellular level, it can be concluded that the main adaptation regarding structure–property relationships toward the optimization of water transport and mechanical stability is mainly achieved at the cell level.
ABSTRACTWe have produced the first annually resolved, centennial‐length tree‐ring chronologies, based on tracheid radial diameter (TRD) and microfibril angle for south‐eastern Australia (SEA) from what would commonly be considered a dendroclimatically suboptimal site. The chronologies exhibit a strong regional temperature signal for the austral summer (nominally November–April) that extends across much of SEA. The strength and spatial extent of the temperature–TRD correlations surpass those between the iconic Tasmanian Mt Read ring‐width chronology and austral summer temperatures, and are more time‐stable. We demonstrate that the value of wood property chronologies for their ability to improve the both the quality and the quantity of highly climate‐sensitive series available for regional annual‐resolution climate reconstructions, in data‐sparse regions in Australasia and beyond, should be examined. In light of the ‘divergence debate’, the time‐stability of relationships with climate, relative to other tree‐ring proxies, also requires further investigation.
In order to investigate the response to climate changes in radial growth of Picea crassifolia at the lower tree line in the middle Qilian mountains in northwestern China, relationships of standardized chronologies of annual ring, earlywood and latewood widths with mean monthly temperature and total monthly precipitation were analyzed by ways of correlation and pointer year analyses. The results show that annual ring, earlywood and latewood widths are significantly negatively correlated with mean monthly temperature in June and July. Annual ring and earlywood widths are significantly and positively correlated with total monthly precipitation in March, May and June and negatively correlated with total monthly precipitation in September. Latewood width is less sensitive to climate changes than the width of earlywood and insignificantly sensitive to precipitation. The results of pointer year analysis revealed that when summer temperatures are higher than the mean summer temperature synchronization and the summer precipitation lower than mean summer precipitation synchronization, narrow annual rings are formed. Wide annual rings are formed when summer temperatures are lower than the mean summer temperature synchronization and summer precipitation higher than mean summer precipitation synchronization. The results indicate that more precipitation in the spring and summer is helpful for radial growth while warmer summer restricts radial growth of P. crassifolia at the lower tree line in the middle Qilian mountains.