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"Xylem - anatomy "
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Atlas of wood, bark and pith anatomy of Eastern Mediterranean trees and shrubs : with a special focus on Cyprus
This atlas presents anatomical descriptions of the xylem, bark and pith of 264 species belonging to 69 families. It highlights the anatomical diversity of trees, shrubs, dwarf shrubs, woody lianas and several of the prominent perennial herbs from the Eastern Mediterranean region, with a focus on the island of Cyprus. The island's topography and biogeographic history combine to provide a wide range of habitats and diverse flora including widespread, endemic, and ornamental species. The monograph for each species includes a description of the anatomical structures of the stem and twig xylem and the twig's bark and pith, as well as color micrographs of double-stained sections of each of these plant parts. These entries are accompanied by a photograph and a brief description of the plant including stem wood density, height, habit, flower, leaf and fruit characteristics, and a map showing its geographic and altitudinal distribution in the region. Xylem descriptions follow the IAWA lists of microscopic features for hardwood and softwood identification. For bark and pith descriptions, a new coding system developed by the authors is applied. Lastly, the atlas offers a key for wood identification that was developed to differentiate between groups of species by using a small number of features that are unambiguous and clearly visible. The atlas will be a valuable guide for botanists, ecologists, foresters, archeologists, horticulturists and paleobotanists.
Book
A broad survey of hydraulic and mechanical safety in the xylem of conifers
Drought-induced forest dieback has been widely reported over the last decades, and the evidence for a direct causal link between survival and hydraulic failure (xylem cavitation) is now well known. Because vulnerability to cavitation is intimately linked to the anatomy of the xylem, the main objective of this study was to better understand the xylem anatomical properties associated with cavitation resistance. An extensive data set of cavitation resistance traits and xylem anatomical properties was developed for 115 conifer species, with special attention given to the micro-morphology of bordered pits. The ratio of torus to pit aperture diameter, so-called torus overlap, increased with increasing cavitation resistance, while the flexibility of the margo does not seem to play a role, suggesting that air-seeding is located at the seal between the aspirated torus and pit aperture. Moreover, punctured tori were reported in various Pinaceae species. Species resistant to cavitation had thicker tracheid walls, while their lumen diameter (conduit size) was only slightly reduced, minimizing the impact on hydraulic conductance. The results also demonstrated (i) the existence of an indirect trade-off between hydraulic safety and mechanical strength; and (ii) a consistency between species distribution and xylem anatomy: species with a wide torus overlap and high valve effects are found in arid environments such as the Mediterranean region.
Journal Article
Conflicting functional effects of xylem pit structure relate to the growth-longevity trade-off in a conifer species
Consistent with a ubiquitous life history trade-off, trees exhibit a negative relationship between growth and longevity both among and within species. However, the mechanistic basis of this life history trade-off is not well understood. In addition to resource allocation conflicts among multiple traits, functional conflicts arising from individual morphological traits may also contribute to life history trade-offs. We hypothesized that conflicting functional effects of xylem structural traits contribute to the growth-longevity trade-off in trees. We tested this hypothesis by examining the extent to which xylem morphological traits (i.e., wood density, tracheid diameters, and pit structure) relate to growth rates and longevity in two natural populations of the conifer species Pinus ponderosa. Hydraulic constraints arise as trees grow larger and xylem anatomical traits adjust to compensate. We disentangled the effects of size through ontogeny in individual trees and growth rates among trees on xylem traits by sampling each tree at multiple trunk diameters. We found that the oldest trees had slower lifetime growth rates compared with younger trees in the studied populations, indicating a growth-longevity trade-off. We further provide evidence that a single xylem trait, pit structure, with conflicting effects on xylem function (hydraulic safety and efficiency) relates to the growth-longevity trade-off in a conifer species. This study highlights that, in addition to trade-offs among multiple traits, functional constraints based on individual morphological traits like that of pit structure provide mechanistic insight into how and when life history trade-offs arise.
Journal Article
Hydraulic adjustment of Scots pine across Europe
The variability of branch-level hydraulic properties was assessed across 12 Scots pine populations covering a wide range of environmental conditions, including some of the southernmost populations of the species. The aims were to relate this variability to differences in climate, and to study the potential tradeoffs between traits. Traits measured included wood density, radial growth, xylem anatomy, sapwood- and leaf-specific hydraulic conductivity (K(S) and K(L)), vulnerability to embolism, leaf-to-sapwood area ratio (A(L) : A(S)), needle carbon isotope discrimination (delta 13C) and nitrogen content, and specific leaf area. Between-population variability was high for most of the hydraulic traits studied, but it was directly associated with climate dryness (defined as a combination of atmospheric moisture demand and availability) only for A(L) : A(S), K(L) and delta 13C. Shoot radial growth and A(L) : A(S) declined with stand development, which is consistent with a strategy to avoid exceedingly low water potentials as tree size increases. In addition, we did not find evidence at the intraspecific level of some associations between hydraulic traits that have been commonly reported across species. The adjustment of Scots pine's hydraulic system to local climatic conditions occurred primarily through modifications of A(L) : A(S) and direct stomatal control, whereas intraspecific variation in vulnerability to embolism and leaf physiology appears to be limited.
Journal Article
Climate of origin shapes variations in wood anatomical properties of 17 Picea species
Background
Variations in hydraulic conductivity may arise from species-specific differences in the anatomical structure and function of the xylem, reflecting a spectrum of plant strategies along a slow-fast resource economy continuum. Spruce (
Picea
spp.), a widely distributed and highly adaptable tree species, is crucial in preventing soil erosion and enabling climate regulation. However, a comprehensive understanding of the variability in anatomical traits of stems and their underlying drivers in the
Picea
genus is currently lacking especially in a common garden.
Results
We assessed 19 stem economic properties and hydraulic characteristics of 17
Picea
species grown in a common garden in Tianshui, Gansu Province, China. Significant interspecific differences in growth and anatomical characteristics were observed among the species. Specifically, xylem hydraulic conductivity (
K
s
) and hydraulic diameter exhibited a significant negative correlation with the thickness to span ratio (TSR), cell wall ratio, and tracheid density and a significant positive correlation with fiber length, and size of the radial tracheid. PCA revealed that the first two axes accounted for 64.40% of the variance, with PC1 reflecting the trade-off between hydraulic efficiency and mechanical support and PC2 representing the trade-off between high embolism resistance and strong pit flexibility. Regression analysis and structural equation modelling further confirmed that tracheid size positively influenced
K
s
, whereas the traits DWT, D_r, and TSR have influenced
K
s
indirectly. All traits failed to show significant phylogenetic associations. Pearson’s correlation analysis demonstrated strong correlations between most traits and longitude, with the notable influence of the mean temperature during the driest quarter, annual precipitation, precipitation during the wettest quarter, and aridity index.
Conclusions
Our results showed that xylem anatomical traits demonstrated considerable variability across phylogenies, consistent with the pattern of parallel sympatric radiation evolution and global diversity in spruce. By integrating the anatomical structure of the stem xylem as well as environmental factors of origin and evolutionary relationships, our findings provide novel insights into the ecological adaptations of the
Picea
genus.
Journal Article
Relationships between xylem vessel characteristics, calculated axial hydraulic conductance and size-controlling capacity of peach rootstocks
Background and Aims Previous studies indicate that the size-controlling capacity of peach rootstocks is associated with reductions of scion water potential during mid-day that are caused by the reduced hydraulic conductance of the rootstock. Thus, shoot growth appears to be reduced by decreases in stem water potential. The aim of this study was to investigate the mechanism of reduced hydraulic conductance in size-controlling peach rootstocks. Methods Anatomical measurements (diameter and frequency) of xylem vessels were determined in shoots, trunks and roots of three contrasting peach rootstocks grown as trees, each with different size-controlling characteristics: ‘Nemaguard’ (vigorous), ‘P30-135’ (intermediate vigour) and ‘K146-43’ (substantially dwarfing). Based on anatomical measurements, the theoretical axial xylem conductance of each tissue type and rootstock genotype was calculated via the Poiseuille–Hagen law. Key Results Larger vessel dimensions were found in the vigorous rootstock (‘Nemaguard’) than in the most dwarfing one (‘K146-43’) whereas vessels of ‘P30-135’ had intermediate dimensions. The density of vessels per xylem area in ‘Nemaguard’ was also less than in ‘P30-135’and ‘K146-43’. These characteristics resulted in different estimated hydraulic conductance among rootstocks: ‘Nemaguard’ had higher theoretical values followed by ‘P30-135’ and ‘K146-43’. Conclusions These data indicate that phenotypic differences in xylem anatomical characteristics of rootstock genotypes appear to influence hydraulic conductance capacity directly, and therefore may be the main determinant of dwarfing in these peach rootstocks.
Journal Article
Modelling the mechanical behaviour of pit membranes in bordered pits with respect to cavitation resistance in angiosperms
Background and AimsVarious correlations have been identified between anatomical features of bordered pits in angiosperm xylem and vulnerability to cavitation, suggesting that the mechanical behaviour of the pits may play a role. Theoretical modelling of the membrane behaviour has been undertaken, but it requires input of parameters at the nanoscale level. However, to date, no experimental data have indicated clearly that pit membranes experience strain at high levels during cavitation events.MethodsTransmission electron microscopy (TEM) was used in order to quantify the pit micromorphology of four tree species that show contrasting differences in vulnerability to cavitation, namely Sorbus aria, Carpinus betulus, Fagus sylvatica and Populus tremula. This allowed anatomical characters to be included in a mechanical model that was based on the Kirchhoff–Love thin plate theory. A mechanistic model was developed that included the geometric features of the pits that could be measured, with the purpose of evaluating the pit membrane strain that results from a pressure difference being applied across the membrane. This approach allowed an assessment to be made of the impact of the geometry of a pit on its mechanical behaviour, and provided an estimate of the impact on air-seeding resistance.Key ResultsThe TEM observations showed evidence of residual strains on the pit membranes, thus demonstrating that this membrane may experience a large degree of strain during cavitation. The mechanical modelling revealed the interspecific variability of the strains experienced by the pit membrane, which varied according to the pit geometry and the pressure experienced. The modelling output combined with the TEM observations suggests that cavitation occurs after the pit membrane has been deflected against the pit border. Interspecific variability of the strains experienced was correlated with vulnerability to cavitation. Assuming that air-seeding occurs at a given pit membrane strain, the pressure predicted by the model to achieve this mechanical state corresponds to experimental values of cavitation sensitivity (P50).ConclusionsThe results provide a functional understanding of the importance of pit geometry and pit membrane structure in air-seeding, and thus in vulnerability to cavitation.
Journal Article
Hydraulic constraints limit height growth in trees at high altitude
• Low temperatures limit the fixation of photosynthates and xylogenesis. Here, we hypothesized that reduced longitudinal growth in trees at high altitude is related to the lower hydraulic efficiency of the transport system. • Apical buds of Norway spruce (Picea abies) trees at high and low elevation were heated during 2006 and 2007. At the end of the experiment, trees were felled. Longitudinal increments and tracheid lumen areas were measured along the stem. Apical hydraulic conductivity (k) was estimated from anatomical data. • Before heating, high-altitude trees showed fewer (P = 0.002) and smaller (P = 0.008) apical conduits, and therefore reduced k (P = 0.016) and stem elongation (P < 0.0001), in comparison with trees at low elevation. After 2 yr of heating, k increased at both high (P = 0.014) and low (P = 0.047) elevation. Only high-altitude trees showed increased stem elongation, which reached the same magnitude as that of controls at low elevation (P = 0.735). Heating around the apical shoots did not appear to induce significant changes in conduit dimension along the rest of the stem. • The total number and size of xylem elements at the stem apex are strongly constrained by low temperatures. Trees at high altitude are therefore prevented from building up an efficient transport system, and their reduced longitudinal growth reflects strong hydraulic limitations.
Journal Article
Are hydraulic patterns of lianas different from trees? New insights from Hedera helix
Lianas form long and flexible but disproportionately narrow stems, and thus require particular strategies to maintain the integrity of xylem water transport and ensure supply to large crown areas. The hydraulic architecture of lianas and the respective within-plant coordination of transport efficiency and safety, and the underlying anatomical variations in xylem, are largely unexplored. We analysed Hedera helix, a liana widespread in European temperate forests, with respect to hydraulic and xylem anatomical variations between the main stem and branches, between juvenile and adult life phases, and along the vertical axis. Main stems were significantly less embolism resistant but exhibited a higher hydraulic conductivity than branches. In branches, the cell turgor loss point of leaves decreased, while the embolism resistance and conductivity of xylem, as well as conduit diameters, increased with height. High water-transport capacities allow ivy to compensate for the small cross-section of stems, while the limited resistance to drought-induced xylem dysfunction of the main stem is probably linked to conservative stomatal regulation. Pronounced differences in xylem anatomy, hydraulic efficiency, and safety between the main stem and branches and along the vertical axis are surprisingly similar to those of self-supporting plants, and indicate that the coordination of carbon and water economies requires similar internal adjustments in tall plants.
Journal Article
Spatial distribution and packing of xylem conduits
Premise of the study: The hydraulic properties of the xylem determine the ability of plants to transport water from the soil to the leaves and to cope with important stress factors such as frost and drought. Hydraulic properties have usually been studied as a function of the anatomy of xylem conduits and their pits, but recent studies have proposed that system-level properties, related to the topology of the xylem network, may also play a role. Here we study how the spatial arrangement of conduits in xylem cross sections affects the relationship between mean conduit lumen area and conduit density (packing function) across species. Methods: Point pattern analysis was used to describe the spatial distribution of xylem conduits in 97 woody species. The effect of conduit aggregation on the packing function was tested using phylogenetic generalized least squares. A hydraulic model with an explicit description of the topology of the xylem network was used to interpret the functional significance of our findings. Key results: The spatial arrangement of conduits affected the packing function across species, so that species with aggregated distributions tended to have lower conduit densities for a given conduit size and lower conduit lumen fractions. According to our modeling results, the higher conduit-to-conduit connectivity of species with aggregated distributions allows them to achieve higher hydraulic conductivity. Species with aggregated conduits, however, pay a cost in terms of increased vulnerability to embolism. Conclusions: The spatial arrangement of conduits affects the fundamental structural and functional attributes of the xylem.
Journal Article