Variation in wood density and anatomy in a widespread mangrove species

Wood density is an important plant trait that influences a range of ecological processes, including resistance to damage and growth rates. Wood density is highly dependent on anatomical characteristics associated with the conductive tissue of trees (xylem and phloem) and the fibre matrix in which they occur. Here, we investigated variation in the wood density of the widespread mangrove species Avicennia marina in the Exmouth Gulf in Western Australia and in the Firth of Thames in New Zealand. We assessed how variation in xylem vessel size, fibre wall thickness and proportion of phloem within the wood contributed to variation in wood density and how these characteristics were linked to growth rates. We found the wood density of A. marina to be higher in Western Australia than in New Zealand and to be higher in taller seaward fringing trees than in scrub trees growing high in the intertidal. At the cellular level, high wood density was associated with large xylem vessels and thick fibre walls. Additionally, wood density increased with decreasing proportions of phloem per growth layer of wood. Tree growth rates were positively correlated with xylem vessel size and wood density. We conclude that A. marina can have large xylem vessel sizes and high growth rates while still maintaining high wood density because of the abundance and thickness of fibres in which vessels are found.     

Impact of extreme drought on tree-ring width and vessel anatomical features of Chukrasia tabularis

Multiple sources of evidence suggest an increasing frequency of extreme climatic events during the past century. In Bangladesh, a country strongly influenced by the South Asian monsoon climate, the years 1999 and 2006 were the most severe droughts among the ten drought events identified over the last four decades. We investigated the impact of these two drought events on radial growth and xylem anatomical features of the brevi-deciduous tree species Chukrasia tabularis in a moist tropical forest in Bangladesh. Tree radial growth declined by 54% during the 1999 and 48.7% during the 2006 droughts, respectively. Among the wood anatomical features, the number of vessels (NV) showed the highest sensitivity to drought, with a 45% decrease in the 1999 drought year, followed by total vessel area (TVA) and mean vessel area (MVA). On the other hand, Vessel density (VD) increased by 13% during the 1999 drought but the increase in VD was very low in the drought year 2006. The decreasing vessel area and increasing vessel density indicate xylem hydraulic adaptation of C. tabularis to minimize drought induced cavitation risk and to avoid hydraulic failure. The significant correlations between the Standardized Precipitation Evapotranspiration Index (SPEI) and time series of tree-ring width and vessel variables imply that decline in radial growth and changes in vessel features in C. tabularis are likely to be caused by drought induced water stress. Our analyses suggest that radial growth and wood anatomical features of C. tabularis are highly sensitive to extreme drought events in South Asian moist tropical forests and can be used to reconstruct past droughts and to model tree response to drought stress under future climate conditions.     

Wood anatomical traits as a measure of plant responses to water availability: invasive Acacia mearnsii De Wild. compared with native tree species in fynbos riparian ecotones, South Africa

Riparian ecotones in the fynbos biome of South Africa are heavily invaded by woody invasive alien species, which are known to reduce water supply to downstream environments. To explore whether variation in species-specific functional traits pertaining to drought-tolerance exist, we investigated wood anatomical traits of key native riparian species and the invasive Acacia mearnsii across different water availability proxies. Wood density, vessel resistance against implosion, vessel lumen diameter and vessel wall thickness were measured. Wood density varied significantly between species, with A. mearnsii having denser wood at sites in rivers with high discharge. As higher wood density is indicative of increased drought tolerance and typical of drier sites, this counter-intuitive finding suggests that increased wood density was more closely related to midday water stress, than streamflow quantity per se. Wood density was positively correlated with vessel resistance against implosion. Higher wood density may also be evidence that A. mearnsii is more resistant against drought-induced cavitation than the studied native species. The observed plastic response of A. mearnsii anatomical traits to variable water availability indicates the ability of this species to persist under various environmental conditions. A possible non-causal relationship between wood anatomy and drought tolerance in these riparian systems is discussed.     

Wood anatomy and vessel characteristics of spiny monkey orange (Strychnos spinosa) in Benin (West Africa)

The response of plant species to varying climate conditions in tropical Africa remains poorly understood but can be assessed using wood anatomical traits. These traits play an important role for the adaptive capacity of a species to environmental stress, since environmental conditions can modify the proportion, size, and morphology of wood anatomical elements. This study reports quantitative data on vessel characteristics of the diffuse porous angiosperm Spiny monkey orange (Strychnos spinosa Lam.) in Benin in tropical West Africa. The vessel-related anatomical traits varied with high amplitude (coefficient of variation CV ˃ 25%) between different sites located in different climate zones. The variability of the traits is higher within one climatic zone than between climatic zones, and even more pronounced within trees. Consequently, the climatic zones have less influence on the studied features than local site conditions. However, the study showed that S. spinosa individuals that have numerous vessels also have a high lumen fraction and total ring area. On the other hand, individuals presenting a high vessel density also display vessels of smaller size. The correlation between vessel number and total ring area on the one hand, and between vessel size and lumen fraction on the other hand are highly significant and positive. In Benin, S. spinosa wood anatomical traits are likely linked to local site factors rather than to regional climatic factors.     

Wood anatomy and wood density in shrubs: Responses to varying aridity along transcontinental transects

Wood density plays a key role in ecological strategies and life history variation in woody plants, but little is known about its anatomical basis in shrubs. We quantified the relationships between wood density, anatomy, and climate in 61 shrub species from eight field sites along latitudinal belts between 31° and 35° in North and South America. Measurements included cell dimensions, transverse areas of each xylem cell type and percentage contact between different cell types and vessels. Wood density was more significantly correlated with precipitation and aridity than with temperature. High wood density was achieved through reductions in cell size and increases in the proportion of wall relative to lumen. Wood density was independent of vessel traits, suggesting that this trait does not impose conduction limitations in shrubs. The proportion of fibers in direct contact with vessels decreased with and was independent of wood density, indicating that the number of fiber-vessel contacts does not explain the previously observed correlation between wood density and implosion resistance. Axial and radial parenchyma each had a significant but opposite association with wood density. Fiber size and wall thickness link wood density, life history, and ecological strategies by controlling the proportion of carbon invested per unit stem volume.     

Temporal annotation of high-resolution intra-annual wood density information of Eucalyptus urophylla and its correlation with hydroclimatic conditions

Three different Eucalyptus urophylla clones grown under two different spacing regimes in an experimental site in the state of São Paulo, Brazil, were analyzed to test effects of clone identity, spacing, cambial age and hydroclimatic conditions on high-resolution intra-annual wood density profiles. Since distinct periodic tree-ring boundaries were not visible on the stem cross-sectional surfaces, finding an alternative method for synchronization of density profiles was crucial for the analysis. The challenge was to generate intra- and inter-tree synchronized density profiles that possess high amplitude variation and low phase variation. Thus, we developed a protocol and workflow of how such high-resolution density profiles can be spatially aligned and temporally annotated to enable correlation analyses between trees and with time series of environmental stimuli. Mean wood density was significantly different between clones, but not between the spacings. Wood density increased significantly with increasing cambial age and decreasing growth rate. Principal component analysis showed that the overall variability in the temporally annotated density profiles is dominated by a highly significant common signal. We found significant negative correlation values for precipitation, indicating that water supply is the main driver of stem growth at the site, and providing evidence for the correctness of the method. The developed workflow can easily be adjusted to the analysis of other intra-annual tree-ring features like anatomical xylem cell traits or isotopic signals in the wood. It has a large potential to be used as a general guideline for the synchronization of intra-annual tree-ring traits, especially when distinct tree-ring boundaries are missing, as it is often the case under tropical climatic conditions. The workflow supports the development of spatially aligned and temporally annotated chronologies under non-annual growth rhythms.     

Relationship between wood anatomy,tree-ring widths and wood density of Pinus sylvestris L. and climate at high latitudes in northern Sweden

In this study, wood anatomy, tree-ring width and wood density of Pinus sylvestris at the northern timberline in Fennoscandia were used to identify relationships among the parameters and to screen them for their climatic signals. Furthermore we investigated the influence of the juvenile wood section for all parameters developed. The measurements of wood anatomy were conducted with confocal laser scanning microscopy (CLSM) while the density profiles were produced using an Itrax MultiScanner. We developed chronologies of ring width, wood density and anatomy for a period between 1940 and 2010. Correlations between wood density and wood anatomy were strong in the latewood part. For some wood anatomy and density chronologies youth trends were found in the juvenile part. Wood density decreased from the pith up to the 9th ring and stabilized afterwards, while cell lumen diameter and lumen area increased simultaneously up to the 15th ring. All chronologies contained strong summer temperature signals. The wood anatomical variables provided additional information about seasonal precipitation which could not be found in wood density and tree-ring widths. Our study confirmed previous results stating that the parameter maximum density contains the strongest climate signal, that is, summer temperatures at the northern timberline. Nevertheless, the intra-annual data on tracheid dimensions showed good potential to supply seasonal climatic information and improve our understanding of climatic effects on tree growth and wood formation.     

CAMBIUM, a process-based model of daily xylem development in Eucalyptus

In hardwoods such as Eucalyptus spp., xylem (wood) is a heterogeneous tissue consisting of multiple cell types. As such, xylem development involves multiple complex interactions. To describe and understand xylem development, and ultimately predict the resultant wood properties, a process-based approach to modelling wood property variation is potentially very useful. In this paper, a new model (CAMBIUM), which incorporates concepts of these processes, is described. CAMBIUM predicts how wood density and fibre and vessel anatomical properties vary from pith-to-bark at a daily time step as a function of changing environmental conditions and a set of simulated physiological processes. Simulations from an existing process-based model of stand development (CABALA) are used as inputs. A key feature of CAMBIUM is a model of the interaction between different xylem cell types. Some weaknesses were identified in the ability of the model to simulate vessel spatial patterns and frequencies, emphasizing the complexities inherent in this aspect of angiosperm wood formation. The model was, however, able to provide realistic estimates of short-term variation and temporal ranges in eucalypt fibre diameter and secondary wall development and wood density.     

Wood anatomy and tree growth covary in riparian ash forests along climatic and ecological gradients

Riparian ash forests subjected to seasonal drought are among the most endangered ecosystems in Europe. They are threatened by climate warming causing aridification and by land-use changes modifying river flow. To assess the impacts of these two stress factors on riparian forests, we studied radial growth and xylem anatomical traits in five narrow-leaved ash (Fraxinus angustifolia) stands across wide climatic and ecological gradients from northern Italy to southern Portugal. Radial growth rates and earlywood hydraulic diameter (Dh) were directly correlated, whilst earlywood vessel density and growth rates were inversely associated. Ash growth positively responded to precipitation. Higher and lower rates of growth increase in response to precipitation were found in dry (annual precipitation 357–750 mm, annual water balance −39 to −48 mm) and wet (annual precipitation 1030 mm, annual water balance 27 mm) sites, respectively. Wet conditions in autumn and winter of the year prior to tree-ring formation lead to larger Dh values, except in the wet site where warmer conditions from prior autumn to current spring were positively associated to wider vessels. Growth was also enhanced by a higher river flow, reflecting higher soil moisture due to elevated groundwater table levels. Peaks in river flow from late winter to early spring increased Dh in dry-continental sites. Growth and potential hydraulic conductivity in drought-prone riparian ash forests are differently impacted by climate variability and river flow depending on site and hydrological conditions. Nevertheless, covariation between radial growth and the earlywood vessel diameter was found, regardless of site specific differences. Wood production and hydraulic conductivity are coordinated through the production of large earlywood vessels which may allow reaching higher growth rates.     

海桑次生木质部导管解剖特征与土壤理化因子年内动态变化的关系研究

为揭示海桑次生木质部导管解剖特征随土壤理化因子年内动态变动而变化的适应机制,该研究利用海桑具有生长轮的特点,通过显微技术界定了海桑采样枝条一年内10个不同连续时间段形成的新"生长层"(新形成的次生木质部),观测了10个不同连续时间段新"生长层"的导管解剖特征,并对10个新"生长层"形成阶段所对应的土壤理化因子数量特征进行了测定,用逐步回归法分析了10个不同连续时间段海桑新"生长层"管孔数量解剖特征与对应土壤理化因子数量特征之间的关系。结果显示:(1)10个不同连续时间段海桑形成新"生长层"管孔数量特征指标,除相邻管孔间接触壁长占比无显著差异外,其他8项指标包括管孔径向直径、管孔弦向直径、导管壁厚、导管长度、管孔密度、单孔率、导管聚合度和相邻管孔间接触壁长等均具有显著差异(P0.05);多重比较显示,10个不同连续时间段海桑形成新"生长层"管孔数量特征8项指标具不同程度的变动。(2)海桑新"生长层"形成阶段(10个不同连续时间段)土壤理化因子,包括土壤有机质含量、土壤全氮含量、土壤全磷含量、土壤pH值和土壤全盐量等5项指标均具有显著差异(P0.05);多重比较显示,海桑新"生长层"形成阶段(10个不同连续时间段)土壤理化因子指标均具有不同程度变动。(3)10个不同连续时间段海桑形成新"生长层"的管孔数量解剖特征,与所对应新"生长层"形成阶段土壤理化因子数量特征的逐步回归分析表明,随着土壤全盐含量的升高,海桑导管弦向直径和导管聚合度同时呈显著增大趋势(P0.05)。研究表明,海桑在一年内不同连续时间段,随土壤全盐量增加,土壤渗透势将增大,水分在次生木质部导管中输导的安全性将下降,而海桑导管分子随一年内不同连续时间段土壤全盐量的增加呈增大趋势,根据木材生态解剖学的观点,水分输导的安全性将进一步降低,但导管聚合度随一年内不同连续时间段土壤全盐量的增加而增大,具有增进水分输导安全性的作用,这可能是海桑对土壤盐含量变化的生态适应策略。     

Biomass dynamics in a logged forest: the role of wood density

Wood density (WD) is believed to be a key trait in driving growth strategies of tropical forest species, and as it entails the amount of mass per volume of wood, it also tends to correlate with forest carbon stocks. Yet there is relatively little information on how interspecific variation in WD correlates with biomass dynamics at the species and population level. We determined changes in biomass in permanent plots in a logged forest in Vietnam from 2004 to 2012, a period representing the last 8 years of a 30 years logging cycle. We measured diameter at breast height (DBH) and estimated aboveground biomass (AGB) growth, mortality, and net AGB increment (the difference between AGB gains and losses through growth and mortality) per species at the individual and population (i.e. corrected for species abundance) level, and correlated these with WD. At the population level, mean net AGB increment rates were 6.47 Mg ha^?1 year^?1 resulting from a mean AGB growth of 8.30 Mg ha^?1 year^?1, AGB recruitment of 0.67 Mg ha^?1 year^?1 and AGB losses through mortality of 2.50 Mg ha^?1 year^?1. Across species there was a negative relationship between WD and mortality rate, WD and DBH growth rate, and a positive relationship between WD and tree standing biomass. Standing biomass in turn was positively related to AGB growth, and net AGB increment both at the individual and population level. Our findings support the view that high wood density species contribute more to total biomass and indirectly to biomass increment than low wood density species in tropical forests. Maintaining high wood density species thus has potential to increase biomass recovery and carbon sequestration after logging.     

Declining hydraulic performances and low carbon investments in tree rings predate Scots pine drought-induced mortality

Key message

The retrospective analysis of wood anatomical features evidences how a long-term deterioration of hydraulic performance and carbon use portend drought-induced mortality in Scots pine.

Abstract

Widespread episodes of drought-induced tree mortality are predicted to become more frequent as climate becomes warmer and drier. Nevertheless, growth trends and their links to changes in wood anatomy before tree dies are still poorly understood. Wood anatomical features provide valuable information that can be extracted to infer the mechanisms leading to tree death. In this study, we characterize drought-induced mortality affecting two Scots pine (Pinus sylvestris) sites (Prades and Arcalís) located in the North Eastern Iberian Peninsula. Co-occurring now-dead and living Scots pine trees were sampled and their wood anatomical features were measured and compared. We aimed to detect differences in anatomical features between living and dead trees, and to infer past physiological performances that might have determined their subsequent death or survival. Now-dead trees showed lower tracheid and resin duct production, and smaller radial lumen diameters than co-occurring living trees. At the more xeric Prades site, these anatomical differences were larger and chronic, i.e. were observed over the three studied decades, whilst they were less pronounced at the other, more mesic Arcalís site, where tree mortality episodes were more recent. This indicates that dead trees’ hydraulic conductivity was severely affected and that carbon investment in xylem formation and resin duct production was constrained prior to tree death. Our findings show that both hydraulic deterioration and low carbon allocation to xylem formation were associated to drought-induced mortality in Scots pine. Nevertheless, the temporal dynamics of these processes differed between populations as a function of site climatic conditions.     

Anatomical characteristics of xylem in tree rings and its relationship with environments

The anatomical traits of xylem are the characteristics of tree rings at the cellular and subcellular scales, and are often reflection of environmental signals. Studying the relationships between anatomical traits of xylem and environmental change not only provide physiological explanations to the statistics in dendroclimatology, but can also provide a new vision for studying the adaptation process and response strategies of tree growth to climate change. In this paper, with the relationships between the anatomical characteristics of xylem in tree-rings (cell chronology) and climate change as a main thread, we first outline the basic principles and mechanisms of wood anatomical features to record environmental signals, and expounded the basic methods involved in the process of xylem anatomy. Secondly, we discuss the relationship between the anatomical features of xylem and climate factors. We then propose the following as possible directions of future research based on the existing knowledge gap in the topical area: (1) to explore the temporal and spatial variations in the anatomical characteristics of xylem in tree-rings along radial and tangential directions and the relationships with environmental changes; (2) to explore the threshold of tree growth response to environmental plasticity and adaptation processes; (3) to assess the synergistic and antagonistic effects as well as the formation mechanisms of climate response among different tree-ring proxies, and to determine the specific roles and contributions of major climatic factors during different periods of tree-ring formation.     

Anatomical Characteristics of Cherry Rootstocks as Possible Preselecting Tools for Prediction of Tree Vigor

An anatomical study of roots and stems of five self-rooted cherry rootstocks with different growth control potentials was performed to compare their structure and xylem anatomy. The aim was to correlate anatomical parameters with rootstock dwarfing potential and theoretical hydraulic conductance (k _h), and to evaluate the potential application of anatomical characteristics in the preselection process for prediction of ultimate tree vigor. One of the mechanisms of water transport efficiency reduction in dwarfing rootstock stems is from the rootstock xylem anatomy. Anatomical parameters of ??Gisela 5?? and ??Mazzard?? were typical for dwarfing and vigorous rootstocks, respectively, and were thus suggested as reference rootstocks. Significantly greater vessel diameter and frequency were found in invigorating and dwarfing rootstocks, respectively. Higher k _h was obtained in roots, compared to stems, due to significantly larger vascular elements. Dwarfing rootstocks had lower k _h due to small vessel lumens and percentage and, to a lesser extent, because of low wood/cortex ratios or percentage of wood. A higher percentage of wood or xylem in cherry roots and stems was not always positively correlated with their conductivity and vigor. Thus, these parameters cannot be reliably used in prediction of the ultimate vigor, although this method was previously suggested for some other fruit tree species. The most reliable anatomical parameters for that purpose proved to be vessel frequency, vessel lumen area, and percentage of vessels on wood cross section. These characteristics could thus be an effective way to estimate dwarfing capacity and could be applied in rootstock selection and breeding programs.     

Anatomical characteristics of xylem in tree rings and its relationship with environments北大核心CSCD

The anatomical traits of xylem are the characteristics of tree rings at the cellular and subcellular scales, and are often reflection of environmental signals. Studying the relationships between anatomical traits of xylem and environmental change not only provide physiological explanations to the statistics in dendroclimatology, but can also provide a new vision for studying the adaptation process and response strategies of tree growth to climate change. In this paper, with the relationships between the anatomical characteristics of xylem in tree-rings (cell chronology) and climate change as a main thread, we first outline the basic principles and mechanisms of wood anatomical features to record environmental signals, and expounded the basic methods involved in the process of xylem anatomy. Secondly, we discuss the relationship between the anatomical features of xylem and climate factors. We then propose the following as possible directions of future research based on the existing knowledge gap in the topical area: (1) to explore the temporal and spatial variations in the anatomical characteristics of xylem in tree-rings along radial and tangential directions and the relationships with environmental changes; (2) to explore the threshold of tree growth response to environmental plasticity and adaptation processes; (3) to assess the synergistic and antagonistic effects as well as the formation mechanisms of climate response among different tree-ring proxies, and to determine the specific roles and contributions of major climatic factors during different periods of tree-ring formation.     

Relationships of growth,stable carbon isotope composition and anatomical properties of leaf and xylem in seven mulberry cultivars: a hint towards drought tolerance

• The fast growth of mulberry depends on high water consumption, but considerable variations in drought tolerance exist across different cultivars. Physiological and anatomical mechanisms are important to plant survival under drought. However, few research efforts have been made to reveal the relationships of these two aspects in relation to drought tolerance.
• In this study, growth rates, leaf functional physiology and anatomical characteristics of leaf and xylem of 1‐year‐old saplings of seven mulberry cultivars at a common garden were compared. Their relationships were also explored.
• Growth, leaf physiology and anatomy were significantly different among the tested cultivars. Foliar stable carbon isotope composition (δ¹³C) was negatively correlated with growth rates, and closely related to several leaf and xylem anatomical traits. Particularly, leaf thickness, predicted hydraulic conductivity and vessel element length jointly contributed 77% of the variability in δ¹³C. Cultivar Wupu had small stomata, intermediate leaf thickness, the smallest hydraulically weighted vessel diameter and highest vessel number, and higher δ¹³C; Yunguo1 had high abaxial stomatal density, low specific leaf area, moderate hydraulic conductivity and δ¹³C; these are beneficial features to reduce leaf water loss and drought‐induced xylem embolism in arid areas. Cultivar Liaolu11 had contrasting physiological and anatomical traits compared with the previous two cultivars, suggesting that it might be sensitive to drought.
• Our findings indicate that growth and δ¹³C are closely associated with both leaf and xylem anatomical characteristics in mulberry, which provides fundamental information to assist evaluation of drought tolerance in mulberry cultivars and in other woody trees.

     

Scalariform-to-simple transition in vessel perforation plates triggered by differences in climate during the evolution of Adoxaceae

Background and Aims Angiosperms with simple vessel perforations have evolved many times independently of species having scalariform perforations, but detailed studies to understand why these transitions in wood evolution have happened are lacking. We focus on the striking difference in wood anatomy between two closely related genera of Adoxaceae, Viburnum and Sambucus, and link the anatomical divergence with climatic and physiological insights.Methods After performing wood anatomical observations, we used a molecular phylogenetic framework to estimate divergence times for 127 Adoxaceae species. The conditions under which the genera diversified were estimated using ancestral area reconstruction and optimization of ancestral climates, and xylem-specific conductivity measurements were performed.Key Results Viburnum, characterized by scalariform vessel perforations (ancestral), diversified earlier than Sambucus, having simple perforations (derived). Ancestral climate reconstruction analyses point to cold temperate preference for Viburnum and warm temperate for Sambucus. This is reflected in the xylem-specific conductivity rates of the co-occurring species investigated, showing that Viburnum lantana has rates much lower than Sambucus nigra.Conclusions The lack of selective pressure for high conductive efficiency during early diversification of Viburnum and the potentially adaptive value of scalariform perforations in frost-prone cold temperate climates have led to retention of the ancestral vessel perforation type, while higher temperatures during early diversification of Sambucus have triggered the evolution of simple vessel perforations, allowing more efficient long-distance water transport.     

Tree growth,cambial phenology,and wood anatomy of limber pine at a Great Basin (USA) mountain observatory

Key message

Anatomical features of Pinus flexilis under warmer and drier conditions along an altitudinal transect revealed a shorter growing season and shifts in the timing of wood formation.

Abstract

Future climate change driven by greenhouse warming is expected to increase both frequency and severity of drought events and heat waves. Possible consequences for forest ecosystems include changes in foundation species and extended die-off phenomena. We investigated tree growth under the set of biotic and abiotic conditions, and their interactions, that are expected in a drier and warmer world using mountain observatories designed to capture elevation gradients in the Great Basin of North America. Stem cambial activity, wood anatomy, and radial growth of limber pine (Pinus flexilis) were examined at two different elevations using automated dendrometers and repeated histological microcores in 2013–2014. Mean annual temperature was 3.7° cooler at the higher site, which received 170 mm year^?1 of precipitation more than the lower site. Mean air temperature thresholds for xylogenesis computed using logistic regression were 7.7 and 12.0 °C at the higher and lower site, respectively. No differences in the onset date of cambial activity were found under such naturally contrasted conditions, with the global change analog provided by the lower site. Growing season was shortened by increasing drought stress at the lower site, thereby reducing xylem production. Stem expansion was only detectable by automated dendrometers at the higher site. Using elevation to simulate climatic changes and their realized ecosystem feedbacks, it was possible to express tree responses in terms of xylem phenology and anatomical adaptations.

     

Living on the edge: Legacy of water availability on Tetraclinis articulata secondary growth under semiarid conditions in Morocco

Tetraclinis articulata is a xerothermic Mediterranean conifer native from NW Africa that grows under semiarid conditions in the Atlas Mountains. This species is particularly well adapted to water scarcity and forms xeric woodlands subjected to recurring drought, aridification and overexploitation. Wood of T. articulata has been considered of limited dendrochronological value due to abundant anatomical anomalies in their growth rings. We studied tree-ring growth and tree-ring features to check its dendrochronological potential and evaluate their climatic signals and drought legacy effects at two semiarid sites with contrasting elevation in Morocco. Tree-ring boundaries were properly identified on wood cores from 60 to 68% of the sampled trees. Intra-annual wood density fluctuations (IADF), microrings, missing rings and rings with undefined limits were most abundant at the low-elevation coastal site than at the mountain site. Microring and missing ring abundance was inversely related to tree-ring growth, independent of cambial age, and mostly dependent on drought occurrence. IADF frequency was independent of tree-ring growth but inversely related to cambial age. A complex response of IADF formation to cool March to June conditions was found at the low-elevation coastal site, while IADF formation in the mountain site was related to May water availability. Undefined ring limits were observed only at the low-elevation site, showing positive relationships with elevated maximum temperatures in previous winter. Previous winter rainfall favored radial growth, but a significant legacy effect of water availability was evident up to three years prior to growth. Tree-ring growth was mostly related to water availability 33 and 26 months prior to growth at low-elevation and mountain sites, respectively. The obtained results reveal a very plastic cambial activity and a strong ability of T. articulata to withstand drought even for long periods.     

Broad Anatomical Variation within a Narrow Wood Density Range—A Study of Twig Wood across 69 Australian Angiosperms

ObjectivesJust as people with the same weight can have different body builds, woods with the same wood density can have different anatomies. Here, our aim was to assess the magnitude of anatomical variation within a restricted range of wood density and explore its potential ecological implications.MethodsTwig wood of 69 angiosperm tree and shrub species was analyzed. Species were selected so that wood density varied within a relatively narrow range (0.38–0.62 g cm^-3). Anatomical traits quantified included wood tissue fractions (fibres, axial parenchyma, ray parenchyma, vessels, and conduits with maximum lumen diameter below 15 μm), vessel properties, and pith area. To search for potential ecological correlates of anatomical variation the species were sampled across rainfall and temperature contrasts, and several other ecologically-relevant traits were measured (plant height, leaf area to sapwood area ratio, and modulus of elasticity).ResultsDespite the limited range in wood density, substantial anatomical variation was observed. Total parenchyma fraction varied from 0.12 to 0.66 and fibre fraction from 0.20 to 0.74, and these two traits were strongly inversely correlated (r = -0.86, P < 0.001). Parenchyma was weakly (0.24 ≤|r|≤ 0.35, P < 0.05) or not associated with vessel properties nor with height, leaf area to sapwood area ratio, and modulus of elasticity (0.24 ≤|r|≤ 0.41, P < 0.05). However, vessel traits were fairly well correlated with height and leaf area to sapwood area ratio (0.47 ≤|r|≤ 0.65, all P < 0.001). Modulus of elasticity was mainly driven by fibre wall plus vessel wall fraction rather than by the parenchyma component.ConclusionsOverall, there seem to be at least three axes of variation in xylem, substantially independent of each other: a wood density spectrum, a fibre-parenchyma spectrum, and a vessel area spectrum. The fibre-parenchyma spectrum does not yet have any clear or convincing ecological interpretation.