Vessel Traits Research Articles

Xylem vessel traits predict the leaf phenology of native and non‐native understorey species of temperate deciduous forests

Summary Non‐native understorey woody species have been shown to extend leaf display and inhabit vacant phenological niches in early spring and late autumn when growing with native counterparts in temperate deciduous forests across the world. Despite the potential competitive advantages, extended leaf duration also subjects non‐native species to possible hydraulic risks associated with maintaining leaves during periods of increased frost probability. It remains unclear how non‐native species are able to maintain xylem function within this context. Leaf phenology in temperate deciduous trees has been shown to be a function of xylem anatomy, with earlier bud break associated with smaller xylem vessels due to the presumed resistance of smaller vessels to freezing‐induced cavitation. We examined relationships between leaf phenology and xylem vessel traits across 82 native and non‐native understorey deciduous woody species common to eastern U.S. deciduous forests. We hypothesized that non‐native species possess xylem vessel traits associated with maximum hydraulic safety during frost‐prone spring and autumn leaf display without compromising rapid growth rate. Larger metaxylem vessels in non‐native species were associated with both faster spring growth and delayed autumn leaf fall compared to native species. Non‐native species also had smaller latewood vessel diameter, latewood vessel area percentage and a higher proportion of solitary vessels in the entire secondary xylem cross section compared to natives, potentially increasing their resistance to freezing‐ and/or drought‐induced cavitation in autumn, thus allowing for delayed autumn leaf fall. Native and non‐native species exhibited similar dates of spring bud break and leaf emergence, consistent with similar xylem vessel size and vessel area percentage within metaxylem and earlywood. Within both groups, species with earlier bud and leaf emergence had a higher total percentage of vessel area within metaxylem and earlywood. This suggests understorey species need sufficient water to support their early spring growth at the risk of freezing‐induced cavitation. Our study suggests xylem vessel properties, along with cross‐sectional spatial xylem vessel distribution, reflect the capacity of non‐native plants to thrive in a new environment and deepen our understanding of the physiological mechanisms of successful invasions of non‐native understorey woody plant species.

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.

Disentangling the effects of ontogeny and environmental factors on xylem anatomy in a semiarid chamaephyte

The hydraulic system of plants reflects a balance between efficiency and safety which is continuously adjusted along plant's life to guarantee survival under changing conditions. One of such adjustments is the formation of new conductive cells, so vessel traits along ring series reflect part of plant's hydraulic plasticity. In this work we disentangled the effects of ontogeny and environmental factors on wood anatomy of a semiarid chamaephyte, Linum suffruticosum. We sampled individuals on contrasting slope aspects in the Ebro valley (Spain), and measured ring width and vessel traits to obtain annually-resolved data series. General Additive Models (GAMs) were used to identify the variability associated to cambial age, slope aspect, and climatic factors: a shift towards a more efficient xylem configuration was observed at the eighth year of life; narrower rings characterized by higher vessel density and smaller vessel sizes were found at south-facing slopes; and low precipitation at the onset of the growing season resulted in safer vessel features. The combination of quantitative analysis of vessels along ring series and GAMs allowed a global understanding of xylem's functional plasticity; moreover, the need for adequately assessing ontogenetic and topographic effects when predicting plant distribution under global change scenarios was reinforced.

Wood specific gravity and anatomy of branches and roots in 113 Amazonian rainforest tree species across environmental gradients

Wood specific gravity (WSG) is a strong predictor of tree performance across environmental gradients. Yet it remains unclear how anatomical elements linked to different wood functions contribute to variation in WSG in branches and roots across tropical forests. We examined WSG and wood anatomy in white sand, clay terra firme and seasonally flooded forests in French Guiana, spanning broad environmental gradients found throughout Amazonia. We measured 15 traits relating to branches and small woody roots in 113 species representing the 15 most abundant species in each habitat and representative species from seven monophyletic lineages occurring in all habitats. Fiber traits appear to be major determinants of WSG, independent of vessel traits, in branches and roots. Fiber traits and branch and root WSG increased from seasonally flooded species to clay terra firme species and lastly to white sand species. Branch and root wood traits were strongly phylogenetically constrained. Lineages differed in wood design, but exhibited similar variation in wood structure across habitats. We conclude that tropical trees can invest differently in support and transport to respond to environmental conditions. Wind disturbance and drought stress represent significant filters driving tree distribution of Amazonian forests; hence we suggest that biophysical explanations should receive more attention.

Are C-Reactive Protein Associated Genetic Variants Associated with Serum Levels and Retinal Markers of Microvascular Pathology in Asian Populations from Singapore?

IntroductionC-reactive protein (CRP) levels are associated with cardiovascular disease and systemic inflammation. We assessed whether CRP-associated loci were associated with serum CRP and retinal markers of microvascular disease, in Asian populations.MethodsGenome-wide association analysis (GWAS) for serum CRP was performed in East-Asian Chinese (N = 2,434) and Malays (N = 2,542) and South-Asian Indians (N = 2,538) from Singapore. Leveraging on GWAS data, we assessed, in silico, association levels among the Singaporean datasets for 22 recently identified CRP-associated loci. At loci where directional inconsistencies were observed, quantification of inter-ethnic linkage disequilibrium (LD) difference was determined. Next, we assessed association for a variant at CRP and retinal vessel traits [central retinal artery equivalent (CRAE) and central retinal vein equivalent (CRVE)] in a total of 24,132 subjects of East-Asian, South-Asian and European ancestry.ResultsSerum CRP was associated with SNPs in/near APOE, CRP, HNF1A and LEPR (p-values ≤4.7×10−8) after meta-analysis of Singaporean populations. Using a candidate-SNP approach, we further replicated SNPs at 4 additional loci that had been recently identified to be associated with serum CRP (IL6R, GCKR, IL6 and IL1F10) (p-values ≤0.009), in the Singaporean datasets. SNPs from these 8 loci explained 4.05% of variance in serum CRP. Two SNPs (rs2847281 and rs6901250) were detected to be significant (p-value ≤0.036) but with opposite effect directions in the Singaporean populations as compared to original European studies. At these loci we did not detect significant inter-population LD differences. We further did not observe a significant association between CRP variant and CRVE or CRAE levels after meta-analysis of all Singaporean and European datasets (p-value >0.058).ConclusionsCommon variants associated with serum CRP, first detected in primarily European studies, are also associated with CRP levels in East-Asian and South-Asian populations. We did not find a causal link between CRP and retinal measures of microvascular disease.

Potential hydraulic efficiency in angiosperm trees increases with growth‐site temperature but has no trade‐off with mechanical strength

AbstractAimXylem structures are closely related to a tree's hydraulic efficiency and mechanical stability, both of which affect the life history and ecological strategy of a species. Although mechanical strength and hydraulic capacity can be shaped by the environment, no such associations between hydraulic efficiency and climatic variables have been reported across a wide range of tree species.LocationYunnan, south‐west China.MethodsWe compiled a data set for vessel density, vessel diameter (D), potential hydraulic conductivity (Kp), wood density (WD), modulus of rupture (MOR) and modulus of elasticity (MOE) from 316 angiosperm tree species. Our objective was to examine the correlations among xylem traits and climatic variables. We hypothesized that both hydraulic efficiency and mechanical strength would vary along climatic gradients, but that a trade‐off would occur between them.ResultsAll xylem traits varied significantly across species, but the magnitudes of variation were greater for vessel traits than for mechanical properties. Values for Kp and D increased with mean annual temperature (MAT) in both evergreen and deciduous trees, but they were significantly correlated with aridity index (AI) in evergreen species only. Both WD and MOR were significantly correlated with MAT only in evergreen trees. MOR decreased with increasing AI in the evergreens, but not in deciduous trees. These findings indicated that xylem development in evergreens is more sensitive to environmental changes than in deciduous trees. However, stem hydraulic traits are independent of mechanical properties.Main conclusionsConsistent with our hypothesis, both hydraulic efficiency and mechanical strength of angiosperm trees are influenced by the environment, with temperature having a more important effect on hydraulic efficiency than precipitation. However, no trade‐off exists between efficiency and strength. This absence of a link is explained because angiosperms have xylem tissue that specifically functions in either mechanical strength or water transport.

Environmental control of vessel traits in Quercus ilex under Mediterranean climate: relating xylem anatomy to function

Time series of tree-ring anatomical features are important to understand the functional role of xylem plasticity over the life span of trees, and thus to reconstruct past ecological and climatic conditions. Holm oak (Quercus ilex L.) is a drought-tolerant tree widely distributed in the Mediterranean Basin. Chronologies of tree-ring width (TRW), vessel lumen area [maximum (MAX) and mean (MVA)] and vessel density (VD) were developed for the period 1942–2001. Each ring was divided into three sections to compare the intra-annual variation of vessel features with the climate conditions during the growing season. The common variability of tree-growth and vessel features was analyzed using a principal component analysis (PCA). Vessel lumen area (MAX and MVA) and TRW loaded positively on the first axis (PC1), whereas VD from the first and second part of the ring (VD1 and VD2) loaded negatively, suggesting that these variables share a common variance. On the other hand, VD in the last third of the ring (VD3) loaded positively on second axis (PC2). PC1 showed a strong positive correlation with precipitation during the hydrological year (prior October–September) and a negative correlation with temperature in spring (April–May), while PC2 showed a negative correlation with precipitation in June. Our results showed that TRW and vessel lumen area were mainly dependent on moisture conditions along the growing season, while vessel density probably plays an important role in the balance between hydraulic conductivity and safety to embolism to better adjust the hydraulic system to water availability.

Wood specific gravity estimation based on wood anatomical traits: Inference of key ecological characteristics in fossil assemblages

Specific gravity (SG) is a key functional trait that has been linked to plant life history traits (e.g. growth rate, life spans) and ecological properties (e.g. cavitation resistance, water transport capacity). SG estimation can thus provide important information about the ecological characteristics of fossil assemblages. Since SG is also related to wood productivity, it can also inform about forest biomass and variation in terrestrial carbon stores over time. Here, we develop a straightforward method to predict SG based on wood anatomical traits that are frequently measured when fossil samples are described. We modeled the relationship of SG as a function of fiber and vessel traits in three extant Mexican communities (135 spp) and selected the best predictive equations. Fiber wall to lumen ratio was the variable most highly related with SG and was present in all the selected prediction models; this variable, together with vessel diameter, was part of the best model. In the validation datasets these models provided good estimates of SG both, at individual sample (overall R2=0.63, varying from 0.44 to 0.84 in different localities) and average community levels. We applied these models to predict SG in three Mexican and one Peruvian fossil assemblages that ranged from Eocene to Miocene in age. To determine the reliability of the estimates, we compared them to observed SG from other extant communities and geographic regions. The estimates of two of the Mexican paleofloras were relatively high, most likely due to deficient preservation. In the other two fossil assemblages, El Cien Formation, Mexico and El Bosque Petrificado Piedra Chamana, Peru, the results were consistent with their inferred paleoenvironment and with the current understanding of the vegetation type they represented. The Peruvian paleoflora had relatively low SG suggesting comparatively higher growth rates and wood productivity, and relatively milder conditions compared to El Cien Formation paleoflora.

Xylem hydraulic adjustment and growth response of Quercus canariensis Willd. to climatic variability

Global change challenges forest adaptability at the distributional limit of species. We studied ring-porous Quercus canariensis Willd. xylem traits to analyze how they adjust to spatio-temporal variability in climate. Trees were sampled along altitudinal transects, and annual time series of radial growth (ring width (RW)) and several earlywood vessel (EV) traits were built to analyze their relationships with climate. The trees responded to increasing water constraints with decreasing altitude and changes in climate in the short term but the analyses showed that xylem did not acclimate in response to long-term temperature increase during the past 30 years. The plants' adjustment to climate variability was expressed in a different but complementary manner by the different xylem traits. At low elevations, trees exhibited higher correlations with water stress indices and trees acclimated to more xeric conditions at low elevations by reducing radial growth and hydraulic diameter (D(H)) but increasing the density of vessels (DV). Average potential conductivity (K(H)) was similar for trees at different altitudes. However, inter-tree differences in xylem traits were higher than those between altitudes, suggesting a strong influence of individual genetic features or micro-site conditions. Trees exhibited higher RW those years with larger D(H) and particularly the linear density of vessels (DV(l)), but partly, climatic signals expressed in RW differed from those in EVs. Trees produced larger D(H) after cold winters and wet years. Ring width responded positively to wet and cool weather in fall and spring, whereas the response to climate of DV and K(H) was generally opposite to that of RW. These relationships likely expressed the negative impact of high respiration rates in winter on the carbon pools used to produce the EVs in the next spring and the overall positive influence of water availability for trees. Our results showed that trees at different sites were able to adjust their hydraulic architecture to climatic variability and temperature increase during recent decades coordinating several complementary traits. Nonetheless, it should be monitored whether they will succeed to acclimate to future climatic scenarios of increasing water stress.

Integration of vessel traits, wood density, and height in angiosperm shrubs and trees

Trees and shrubs tend to occupy different niches within and across ecosystems; therefore, traits related to their resource use and life history are expected to differ. Here we analyzed how growth form is related to variation in integration among vessel traits, wood density, and height. We also considered the ecological and evolutionary consequences of such differences. In a sample of 200 woody plant species (65 shrubs and 135 trees) from Argentina, Mexico, and the United States, standardized major axis (SMA) regression, correlation analyses, and ANOVA were used to determine whether relationships among traits differed between growth forms. The influence of phylogenetic relationships was examined with a phylogenetic ANOVA and phylogenetically independent contrasts (PICs). A principal component analysis was conducted to determine whether trees and shrubs occupy different portions of multivariate trait space. Wood density did not differ between shrubs and trees, but there were significant differences in vessel diameter, vessel density, theoretical conductivity, and as expected, height. In addition, relationships between vessel traits and wood density differed between growth forms. Trees showed coordination among vessel traits, wood density, and height, but in shrubs, wood density and vessel traits were independent. These results hold when phylogenetic relationships were considered. In the multivariate analyses, these differences translated as significantly different positions in multivariate trait space occupied by shrubs and trees. Differences in trait integration between growth forms suggest that evolution of growth form in some lineages might be associated with the degree of trait interrelation.

The Relationship between Retinal Arteriolar and Venular Calibers Is Genetically Mediated, and Each Is Associated with Risk of Cardiovascular Disease

Retinal arteriolar and venular calibers are highly heritable and associated with cardiovascular disease. This study was designed to investigate the relative influence of genetic and environmental factors on the high phenotypic correlation (r = 0.59) between these two traits and to assess the shared and specific influence of established and novel cardiovascular disease risk factors on them. A total of 1463 Caucasian female twins (706 monozygotic and 757 dizygotic), between 24 and 79 years of age, underwent retinal photography from which retinal arteriolar (mean, 153.75 ± 22.1 μm, SD) and venular (mean, 232.1 ± 36.6 μm) calibers were measured with semiautomated software. A bivariate heritability model was used to assess the genetic and environmental influences underlying both specific trait variance and the covariance between the vessel traits. The investigation was an assessment of phenotypic associations between retinal arteriolar and venular calibers and cardiovascular disease risk factors. Additive genetic factors accounted for approximately three fourths of the covariance between retinal arteriolar and venular calibers within the cohort. This finding was replicated in a sample of 1981 twins from the Australian Twins Eye Study. The partial correlation showed that known risk factors accounted for only 5% of the covariance between arteriolar and venular calibers. Novel associations were found between venular caliber and β-cell function (P = 0.011) and insulin sensitivity (P = 0.002). These results suggest that future gene-mapping studies may identify pleiotropic genetic variants influencing both retinal arteriolar and venular calibers. Genetic variants associated with retinal caliber and (risk factors for) cardiovascular disease should provide new etiologic insights into this complex disease.

The importance of wood traits and hydraulic conductance for the performance and life history strategies of 42 rainforest tree species

*In a comparative study of 42 rainforest tree species we examined relationships amongst wood traits, diameter growth and survival of large trees in the field, and shade tolerance and adult stature of the species. *The species show two orthogonal axes of trait variation: a primary axis related to the vessel size-number trade-off (reflecting investment in hydraulic conductance vs hydraulic safety) and a secondary axis related to investment in parenchyma vs fibres (storage vs strength). Across species, growth rate was positively related to vessel diameter and potential specific hydraulic conductance (K(p)), and negatively related to wood density. Survival rate was only positively related to wood density. *Light-demanding species were characterized by low wood and vessel density and wide vessels. Tall species were characterized by wide vessels with low density and large K(p). Hydraulic traits were more closely associated with adult stature than with light demand, possibly because tall canopy species experience more drought stress and face a higher cavitation risk. *Vessel traits affect growth and wood density affects growth and survival of large trees in the field. Vessel traits and wood density are therefore important components of the performance and life history strategies of tropical tree species.

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.

Detecting invisible growth rings of trees in seasonally dry forests in Thailand: isotopic and wood anatomical approaches

We measured radial variation of carbon isotope composition and vessel traits in tree species in seasonally dry forests of Northeast Thailand to explore a more reliable and amenable method of tropical dendrochronology for trees that lack visually detectable and consistent growth rings. Six Dipterocarpaceae species (3 Shorea, 2 Dipterocarpus, and 1 Hopea species) with indistinct or irregular growth rings and teak (Tectona grandis), a species which forms distinct growth rings, were examined. The δ13C value variations in all species showed annual cyclicity. Dipterocarpaceae species usually marked the lowest values of δ13C in the middle of the growing season, whereas teak had the lowest values at nearly the end of the growing season. Since the growing season of the species examined almost corresponds to the rainy season in the study area, the δ13C variation was likely caused by the change in moisture availability. The different variation pattern of teak was attributable to its stronger dependence on 13C-enriched reserved material early in the growing season. Changes in tree vessel traits for all species examined also showed annual cyclicity. Dipterocarpaceae species showed significant correlation between δ13C values and vessel measurements. Vessel lumen (mean area, tangential and radial diameter, and proportion of total area) had a negative correlation, whereas vessel frequency showed a positive correlation. The correlations indicated that changes in vessel traits were caused by the seasonal variation of moisture available to the trees. Thus, we concluded that methods using wood anatomy, as well as δ13C, have great potential for use as tools in tropical dendrochronology within the context of seasonal climate.

Wood density and vessel traits as distinct correlates of ecological strategy in 51 California coast range angiosperms

Wood density and vessel characteristics are functionally interrelated, yet they may have distinct ecological associations. In a comparative study of 51 angiosperm species ranging from chaparral shrubs to riparian trees, we examined relationships among wood density and vessel traits and their ecological correlates. Mean vessel lumen area and vessel density (number mm(-2)) varied widely (7- to 10-fold). In multivariate analyses, both vessel traits were negatively correlated with wood density, which varied more narrowly (< 2-fold). Vessel density and lumen area were inversely related across species, allowing a broad range of vessel traits within a narrow range of wood density. Phylogenetic independent contrasts indicated correlated inverse evolutionary change in vessel traits. Each trait had a distinct pattern of ecological correlation -- wood density was most strongly associated with soil water, and vessel traits showed contrasting relationships with plant height. Within a narrow range of wood density, there was significant variation in vessel traits. Given their particular ecological associations, the results suggest that wood density and vessel traits describe two distinct ecological axes.