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 (Ks) 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 Ks, whereas the traits DWT, D_r, and TSR have influenced Ks 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.

Comparative venation costs of monocotyledon and dicotyledon species in the eastern Colorado steppe.

MAIN CONCLUSION: Leaf vein network cost (total vein surface area per leaf volume) for major veins and vascular bundles did not differ between monocot and dicot species in 21 species from the eastern Colorado steppe. Dicots possessed significantly larger minor vein networks than monocots. Across the tree of life, there is evidence that dendritic vascular transport networks are optimized, balancing maximum speed and integrity of resource delivery with minimal resource investment in transport and infrastructure. Monocot venation, however, is not dendritic, and remains parallel down to the smallest vein orders with no space-filling capillary networks. Given this departure from the "optimized" dendritic network, one would assume that monocots are operating at a significant energetic disadvantage. In this study, we investigate whether monocot venation networks bear significantly greater carbon/construction costs per leaf volume than co-occurring dicots in the same ecosystem, and if so, what physiological or ecological advantage the monocot life form possesses to compensate for this deficit. Given that venation networks could also be optimized for leaf mechanical support or provide herbivory defense, we measured the vascular system of both monocot and dicots at three scales to distinguish between leaf investment in mechanical support (macroscopic vein), total transport and capacitance (vascular bundle), or exclusively water transport (xylem) for both parallel and dendritic venation networks. We observed that vein network cost (total vein surface area per leaf volume) for major veins and vascular bundles was not significantly different between monocot species and dicot species. Dicots, however, possess significantly larger minor vein networks than monocots. The 19 species subjected to gas-exchange measurement in the field displayed a broad range of Amax and but demonstrated no significant relationships with any metric of vascular network size in major or minor vein classes. Given that monocots do not seem to display any leaf hydraulic disadvantage relative to dicots, it remains an important research question why parallel venation (truly parallel, down to the smallest vessels) has not arisen more than once in the history of plant evolution.

Root topological order drives variation of fine root vessel traits and hydraulic strategies in tropical trees.

Vessel traits contribute to plant water transport from roots to leaves and thereby influence how plants respond to soil water availability, but the sources of variation in fine root anatomical traits remain poorly understood. Here, we explore the variations of fine root vessel traits along topological orders within and across tropical tree species. Anatomical traits were measured along five root topological orders in 80 individual trees of 20 species from a tropical forest in southwestern China. We found large variations for most root anatomical traits across topological orders, and strong co-variations between vessel traits. Within species, theoretical specific xylem hydraulic conductivity (Kth) increased with topological order due to increased mean vessel diameter, size heterogeneity, and decreased vessel density. Across species, Kth was associated with vessel fraction in low-order roots and correlated with mean vessel diameter and vessel density in high-order roots, suggesting a shift in relative anatomical contributors to Kth from the second- to fifth-order roots. We found no clear relationship between Kth and stele: root diameter ratios. Our study shows strong variations in root vessel traits across topological orders and species, and highlights shifts in the anatomical underpinnings by varying vessel-related anatomical structures for an optimized water supply.

Modeling Xylem Functionality Aspects.

Depending on the questions to be answered, water flow in the xylem can be modelled following different approaches with varying spatial and temporal resolution. When focussing on the influence of hydraulic architecture upon flow dynamics, distribution of water potentials in a tree crown or questions of vulnerability of the hydraulic system, functional-structural plant models, which link representations of morphological structure with simulated processes and with a virtual environment, can be a promising tool. Such a model will then include a network of idealized xylem segments, each representing the conducting part of a stem or branch segment, and a numerical machinery suitable for solving a system of differential equations on it reflecting the hydrodynamic laws, which are the basis of the broadly accepted cohesion-tension theory of water flow in plants. We will discuss functional-structural plant models, the simplifications that are useful for hydraulic simulations within this framework, the deduction of the used differential equations from basic physical conservation laws, and their numerical solution, as well as additional necessary models of radiation, photosynthesis, and stomatal conductance. In some supplementary notes, we are shortly addressing some related questions, for example, about root systems or about the relation between macro-scale hydraulic parameters and fine-grained (anatomical) xylem structure.

Do root secondary xylem functional traits differ between growth forms in Fabaceae species in a seasonally dry Neotropical environment?

BACKGROUND AND AIMS: Whole-plant performance in water-stressed and disturbance-prone environments depends on a suitable supply of water from the roots to the leaves, storage of reserves during periods of shortage, and a morphological arrangement that guarantees the maintenance of the plants anchored to the soil. All these functions are performed by the secondary xylem of roots. Here, we investigate whether different growth forms of Fabaceae species from the seasonally dry Neotropical environment have distinct strategies for water transport, mechanical support and non-structural carbon and water storage in the root secondary xylem. METHODS: We evaluated cross-sections of root secondary xylem from species of trees, shrubs and subshrubs. We applied linear models to verify the variability in secondary xylem anatomical traits among growth forms. KEY RESULTS: Secondary xylem with larger vessels and lower vessel density was observed in tree species. Vessel wall thickness, vessel grouping index, potential hydraulic conductivity and cell fractions (vessels, fibres, rays and axial parenchyma) were not statistically different between growth forms, owing to the high interspecific variation within the groups studied. CONCLUSION: Our results showed that the variability in anatomical traits of the secondary xylem of the root is species specific. In summary, the cellular complexity of the secondary xylem ensures multiple functional strategies in species with distinct growth forms, a key trait for resource use in an environment with strong water seasonality.

Toward a better understanding of angiosperm xylogenesis: a new method for a cellular approach.

The kinetics of wood formation in angiosperms are largely unknown because their complex xylem anatomy precludes using the radial position of vessels and fibers to infer their time of differentiation. We analyzed xylogenesis in ring-porous ash (Fraxinus angustifolia) and diffuse-porous beech (Fagus sylvatica) over 1 yr and proposed a novel procedure to assess the period of vessel and fiber enlargement using a referential radial file (RRF). Our approach captured the dynamics of wood formation and provided a robust estimation of the kinetics of vessel and fiber enlargement. In beech, fibers and vessels had a similar duration of enlargement, decreasing from 14 to 5 d between April and July. In ash, wide vessels formed in April enlarged at a rate of 27 × 103 µm2 d-1 , requiring half the time of contemporary fibers (6 vs 12 d), and less time than the narrower vessels (14 d) formed in May. These findings reveal distinct cell-type-dependent mechanisms for differentiation in diffuse-porous and ring-porous trees, enhancing our understanding of angiosperm wood cell kinetics. Our approach presents an effective method for investigating angiosperm wood formation and provides a more accurate representation of vessel and fiber morphogenesis in wood formation models.

Xylem resistance to cavitation increases during summer in Pinus halepensis.

Cavitation resistance has often been viewed as a relatively static trait, especially for stems of forest trees. Meanwhile, other hydraulic traits, such as turgor loss point (Ψtlp ) and xylem anatomy, change during the season. In this study, we hypothesized that cavitation resistance is also dynamic, changing in coordination with Ψtlp . We began with a comparison of optical vulnerability (OV), microcomputed tomography (µCT) and cavitron methods. All three methods significantly differed in the slope of the curve,Ψ12 and Ψ88 , but not in Ψ50 (xylem pressures that cause 12%, 88%, 50% cavitation, respectively). Thus, we followed the seasonal dynamics (across 2 years) of Ψ50 in Pinus halepensis under Mediterranean climate using the OV method. We found that Ψ50 is a plastic trait with a reduction of approximately 1 MPa from the end of the wet season to the end of the dry season, in coordination with the dynamics of the midday xylem water potential (Ψmidday ) and the Ψtlp . The observed plasticity enabled the trees to maintain a stable positive hydraulic safety margin and avoid cavitation during the long dry season. Seasonal plasticity is vital for understanding the actual risk of cavitation to plants and for modeling species' ability to tolerate harsh environments.

Study of the mechanism of embolism removal in xylem vessels by using microfluidic devices.

Determining the mechanism that effects embolism repair in the xylem vessels of plants is of great significance in predicting plant distribution and the screening of drought-resistant plants. However, the mechanism of perforation plates of xylem vessels in the acceleration of embolism repair is still not clear by using conventional methods of anatomy and visualization technology. Microfluidic devices have shown their ability to simulate physiological environments and conduct quantitative experiments. This work proposes a biomimetic microfluidic device to study the mechanism of perforation plates of xylem vessels in the acceleration of embolism repair. The results proffered that the perforation plates increase the rate of embolism removal by increasing the pressure differential through the vessel, and the rate of embolism removal is related to the structural parameters of the perforation plate. A combination of the perforation size, the vessel diameter and the perforation plate angle can be optimised to generate higher pressure differentials, which can accelerate the process of embolism repair. This work provides a new method for studying the mechanism of microstructures of natural plants. Furthermore, the mechanism that perforation plates accelerate embolism repair was applied to an electrochemical flow sensor for online determination of heavy metal ions. Test results of this application indicate that the mechanism can be applied in the engineering field to solve the problems of reduced sensitivity of devices, inaccuracy of analysis results and poor reaction performance caused by bubbles that are generated or introduced easily in microdevices, which paves the way for applying the theory to engineering.

Addressing controversies in the xylem embolism resistance-vessel diameter relationship.

Although xylem embolism is a key process during drought-induced tree mortality, its relationship to wood anatomy remains debated. While the functional link between bordered pits and embolism resistance is known, there is no direct, mechanistic explanation for the traditional assumption that wider vessels are more vulnerable than narrow ones. We used data from 20 temperate broad-leaved tree species to study the inter- and intraspecific relationship of water potential at 50% loss of conductivity (P50 ) with hydraulically weighted vessel diameter (Dh ) and tested its link to pit membrane thickness (TPM ) and specific conductivity (Ks ) on species level. Embolism-resistant species had thick pit membranes and narrow vessels. While Dh was weakly associated with TPM , the P50 -Dh relationship remained highly significant after accounting for TPM . The interspecific pattern between P50 and Dh was mirrored by a link between P50 and Ks , but there was no evidence for an intraspecific relationship. Our results provide robust evidence for an interspecific P50 -Dh relationship across our species. As a potential cause for the inconsistencies in published P50 -Dh relationships, our analysis suggests differences in the range of trait values covered, and the level of data aggregation (species, tree or sample level) studied.

A safe breeding ground: genetically improved maritime pine for growth and stem form has more efficient but not more vulnerable xylem.

Through repeated cycles of selection and recombination, tree breeding programs deliver genetically improved varieties for a range of target characteristics such as biomass production, stem form, resistance to biotic stresses, wood properties, etc. However, in the context of increased drought and heat waves, it is not yet known whether growth performance will impede drought resistance. To address this question, we compared the hydraulic properties, such as hydraulic efficiency and hydraulic safety, in four varieties over successive varieties of genetically improved maritime pines (i.e., Pinus pinaster Aït.) for growth and stem form. We measured 22 functional traits related to hydraulic efficiency, hydraulic safety, xylem anatomy and wood density. We found that improved varieties presented higher hydraulic conductivity with larger tracheid lumen size and tracheid lumen fraction, and smaller wall thickness reinforcement and tracheid density, but not at the cost of reduced embolism resistance. The reported absence of trade-off between hydraulic conductivity and embolism resistance is a strong asset to improve biomass productivity, through increased hydraulic efficiency, without impacting drought resistance, and should enable new maritime pine varieties to cope with a drier climate. Our study is one of the first to reveal the hydraulic mechanisms over successive varieties of genetic improvement for tree growth. It provides guidelines for sustainable forest management through breeding for other forest tree species.

A protoxylem pathway to evolution of pith? An hypothesis based on the Early Devonian euphyllophyte Leptocentroxyla.

BACKGROUND AND AIMS: The Early Devonian (Emsian, 400-395 Ma) tracheophyte Leptocentroxyla tetrarcha Bickner et Tomescu emend. Tomescu et McQueen combines plesiomorphic Psilophyton-type tracheid thickenings with xylem architecture intermediate between the plesiomorphic basal euphyllophyte haplosteles and the complex actinosteles of Middle Devonian euphyllophytes. We document xylem development in Leptocentroxyla based on anatomy and explore its implications, which may provide a window into the evolution of pith. METHODS: Leptocentroxyla is preserved by permineralization in the Battery Point Formation (Quebec, Canada). Serial sections obtained using the cellulose acetate peel technique document branching pattern, anatomy of trace divergence to appendages, protoxylem architecture, and variations in tracheid size and wall thickening patterns. KEY RESULTS: Leptocentroxyla has opposite decussate pseudo-whorled branching and mesarch protoxylem, and represents the earliest instance of central histological differentiation in a euphyllophyte actinostele. Tracheids at the centre of xylem exhibit simplified Psilophyton-type wall thickenings and are similar in size (at the axis centre) or smaller than the surrounding metaxylem tracheids (at the centre of appendage traces). CONCLUSIONS: The position and developmental attributes of the simplified Psilophyton-type tracheids suggest they may have been generated by the protoxylem developmental pathway. This supports the delayed and shortened protoxylem differentiation hypothesis, which explains the evolution of pith by (1) delay in the onset of differentiation and lengthening of cell growth duration in a central protoxylem strand; and (2) shortening of the interval of differentiation of those tracheids, leading to progressive simplification (and eventual loss) of secondary wall thickenings, and replacement of tracheids with a central parenchymatous area. NAC domain transcription factors and their interactions with abscisic acid may have provided the regulatory substrate for the developmental changes that led to the evolution of pith. These could have been orchestrated by selective pressures associated with the expansion of early vascular plants into water-stresses upland environments.

Diameters of phloem sieve elements can predict stem growth rates of woody plants.

Understanding forest dynamics is crucial to addressing climate change and reforestation challenges. Plant anatomy can help predict growth rates of woody plants, contributing key information on forest dynamics. Although features of the water-transport system (xylem) have long been used to predict plant growth, the potential contribution of carbon-transporting tissue (phloem) remains virtually unexplored. Here, we use data from 347 woody plant species to investigate whether species-specific stem diameter growth rates can be predicted by the diameter of both the xylem and phloem conducting cells when corrected for phylogenetic relatedness. We found positive correlations between growth rate, phloem sieve element diameter and xylem vessel diameter in liana species sampled in the field. Moreover, we obtained similar results for data extracted from the Xylem Database, an online repository of functional, anatomical and image data for woody plant species. Information from this database confirmed the correlation of sieve element diameter and growth rate across woody plants of various growth forms. Furthermore, we used data subsets to explore potential influences of biomes, growth forms and botanical family association. Subsequently, we combined anatomical and geoclimatic data to train an artificial neural network to predict growth rates. Our results demonstrate that sugar transport architecture is associated with growth rate to a similar degree as water-transport architecture. Furthermore, our results illustrate the potential value of artificial neural networks for modeling plant growth under future climatic scenarios.

Comparative anatomy of leaf petioles in temperate trees and shrubs: the role of plant size, environment and phylogeny.

BACKGROUND AND AIMS: Petioles are important plant organs connecting stems with leaf blades and affecting light-harvesting ability of the leaf as well as transport of water, nutrients and biochemical signals. Despite the high diversity in petiole size, shape and anatomy, little information is available regarding their structural adaptations across evolutionary lineages and environmental conditions. To fill this knowledge gap, we investigated the variation of petiole morphology and anatomy of mainly European woody species to better understand the drivers of internal and external constraints in an evolutionary context. METHODS: We studied how petiole anatomical features differed according to whole-plant size, leaf traits, thermal and hydrological conditions, and taxonomic origin in 95 shrubs and trees using phylogenetic distance-based generalized least squares models. KEY RESULTS: Two major axes of variation were related to leaf area and plant size. Larger and softer leaves are found in taller trees of more productive habitats. Their petioles are longer, with a circular outline and are anatomically characterized by the predominance of sclerenchyma, larger vessels, interfascicular areas with fibres and indistinct phloem rays. In contrast, smaller and tougher leaves are found in shorter trees and shrubs of colder or drier habitats. Their petioles have a terete outline, phloem composed of small cells and radially arranged vessels, fibreless xylem and lamellar collenchyma. Individual anatomical traits were linked to different internal and external drivers. Petiole length and vessel diameter increase with increasing leaf blade area. Collenchyma becomes absent with increasing temperature, and petiole outline becomes polygonal with increasing precipitation. CONCLUSIONS: We conclude that species' temperature and precipitation optima, plant height, and leaf area and thickness exerted a significant control on petiole anatomical and morphological structures not confounded by phylogenetic inertia. Species with different evolutionary histories but similar thermal and hydrological requirements have converged to similar petiole anatomical structures.

Convergent tip-to-base widening of water-conducting conduits in the tallest bryophytes.

PREMISE: Tip-to-base conduit widening is considered a key mechanism that enables vascular plants to grow tall by decreasing the hydraulic resistance imposed by increasing height. Widening of hydraulic anatomy (larger conducting elements toward the base of the vascular system) minimizes gradients in leaf-specific hydraulic conductance with plant height, allowing uniform photosynthesis across the crown of trees. Tip-to-base conduit widening has also been associated with changes in conduit number. However, in bryophytes, despite having representatives with internal water-conducting tissue, conduit widening has been scarcely investigated. METHODS: Here, we examined the changes in hydroid diameter and number with distance from plant tip in Dawsonia superba and D. polytrichoides, two representatives of the genus containing the tallest extant bryophytes. RESULTS: The position of these moss species on the global scale of conduit size and plant size was consistent with a general scaling among plants with internal water transport. Within plants, patterns of conduit widening and number with distance from plant tip in endohydric mosses were similar to those observed in vascular plants. CONCLUSIONS: This study demonstrated that land plants growing upward in the atmosphere show analogous conduit widening of hydraulic structures, suggesting that efficient internal water transport is a convergent adaptation for photosynthesis on land.

Pit and tracheid anatomy explain hydraulic safety but not hydraulic efficiency of 28 conifer species.

Conifers face increased drought mortality risks because of drought-induced embolism in their vascular system. Variation in embolism resistance may result from species differences in pit structure and function, as pits control the air seeding between water-transporting conduits. This study quantifies variation in embolism resistance and hydraulic conductivity for 28 conifer species grown in a 50-year-old common garden experiment and assesses the underlying mechanisms. Conifer species with a small pit aperture, high pit aperture resistance, and large valve effect were more resistant to embolism, as they all may reduce air seeding. Surprisingly, hydraulic conductivity was only negatively correlated with tracheid cell wall thickness. Embolism resistance and its underlying pit traits related to pit size and sealing were more strongly phylogenetically controlled than hydraulic conductivity and anatomical tracheid traits. Conifers differed in hydraulic safety and hydraulic efficiency, but there was no trade-off between safety and efficiency because they are driven by different xylem anatomical traits that are under different phylogenetic control.

Alpine dwarf shrubs show high proportions of nonfunctional xylem: Visualization and quantification of species-specific patterns.

Xylem conductive capacity is a key determinant of plant hydraulic function and intimately linked to photosynthesis and productivity, but can be impeded by temporary or permanent conduit dysfunctions. Here we show that persistent xylem dysfunctions in unstressed plants are frequent in Alpine dwarf shrubs and occur in various but species-specific cross-sectional patterns. Combined synchrotron micro-computed tomography (micro-CT) imaging, xylem staining, and flow measurements in saturated samples of six widespread Ericaceae species evidence a high proportion (19%-50%) of hydraulically nonfunctional xylem areas in the absence of drought stress, with regular distribution of dysfunctions between or within growth rings. Dysfunctions were only partly reversible and reduced the specific hydraulic conductivity to 1.38 to 3.57 ×10-4 m2 s-1 MPa-1 . Decommission of inner growth rings was clearly related to stem age and a higher vulnerability to cavitation of older rings, while the high proportion of nonfunctional conduits in each annual ring needs further investigations. The lower the xylem fraction contributing to the transport function, the higher was the hydraulic efficiency of conducting xylem areas. Improved understanding of the functional lifespan of xylem elements and the prevalence and nature of dysfunctions is critical to correctly assess structure-function relationships and whole-plant hydraulic strategies.

Limited plasticity of anatomical and hydraulic traits in aspen trees under elevated CO2 and seasonal drought.

The timing of abiotic stress elicitors on wood formation largely affects xylem traits that determine xylem efficiency and vulnerability. Nonetheless, seasonal variability of elevated CO2 (eCO2) effects on tree functioning under drought remains largely unknown. To address this knowledge gap, 1-year-old aspen (Populus tremula L.) trees were grown under ambient (±445 ppm) and elevated (±700 ppm) CO2 and exposed to an early (spring/summer 2019) or late (summer/autumn 2018) season drought event. Stomatal conductance and stem shrinkage were monitored in vivo as xylem water potential decreased. Additional trees were harvested for characterization of wood anatomical traits and to determine vulnerability and desorption curves via bench dehydration. The abundance of narrow vessels decreased under eCO2 only during the early season. At this time, xylem vulnerability to embolism formation and hydraulic capacitance during severe drought increased under eCO2. Contrastingly, stomatal closure was delayed during the late season, while hydraulic vulnerability and capacitance remained unaffected under eCO2. Independently of the CO2 treatment, elastic, and inelastic water pools depleted simultaneously after 50% of complete stomatal closure. Our results suggest that the effect of eCO2 on drought physiology and wood traits are small and variable during the growing season and question a sequential capacitive water release from elastic and inelastic pools as drought proceeds.

The Widened Pipe Model of plant hydraulic evolution.

Shaping global water and carbon cycles, plants lift water from roots to leaves through xylem conduits. The importance of xylem water conduction makes it crucial to understand how natural selection deploys conduit diameters within and across plants. Wider conduits transport more water but are likely more vulnerable to conduction-blocking gas embolisms and cost more for a plant to build, a tension necessarily shaping xylem conduit diameters along plant stems. We build on this expectation to present the Widened Pipe Model (WPM) of plant hydraulic evolution, testing it against a global dataset. The WPM predicts that xylem conduits should be narrowest at the stem tips, widening quickly before plateauing toward the stem base. This universal profile emerges from Pareto modeling of a trade-off between just two competing vectors of natural selection: one favoring rapid widening of conduits tip to base, minimizing hydraulic resistance, and another favoring slow widening of conduits, minimizing carbon cost and embolism risk. Our data spanning terrestrial plant orders, life forms, habitats, and sizes conform closely to WPM predictions. The WPM highlights carbon economy as a powerful vector of natural selection shaping plant function. It further implies that factors that cause resistance in plant conductive systems, such as conduit pit membrane resistance, should scale in exact harmony with tip-to-base conduit widening. Furthermore, the WPM implies that alterations in the environments of individual plants should lead to changes in plant height, for example, shedding terminal branches and resprouting at lower height under drier climates, thus achieving narrower and potentially more embolism-resistant conduits.

Cavitation fatigue in conifers: a study on eight European species.

After drought-induced embolism and repair, tree xylem may be weakened against future drought events (cavitation fatigue). As there are few data on cavitation fatigue in conifers available, we quantified vulnerability curves (VCs) after embolism/repair cycles on eight European conifer species. We induced 50% and 100% loss of conductivity (LC) with a cavitron, and analyzed VCs. Embolism repair was obtained by vacuum infiltration. All species demonstrated complete embolism repair and a lack of any cavitation fatigue after 50% LC . After 100% LC, European larch (Larix decidua), stone pine (Pinus cembra), Norway spruce (Picea abies), and silver fir (Abies alba) remained unaffected, while mountain pine (Pinus mugo), yew (Taxus baccata), and common juniper (Juniperus communis) exhibited 0.4-0.9 MPa higher vulnerability to embolism. A small cavitation fatigue observed in Scots pine (Pinus sylvestris) was probably biased by incomplete embolism repair, as indicated by a correlation of vulnerability shifts and conductivity restoration. Our data demonstrate that cavitation fatigue in conifers is species-specific and depends on the intensity of preceding LC. The lack of fatigue effects after moderate LC, and relevant effects in only three species after high LC, indicate that conifers are relatively resistant against cavitation fatigue. This is remarkable considering the complex and delicate conifer pit architecture and may be important considering climate change projections.

Developmental and biophysical determinants of grass leaf size worldwide.

One of the most notable ecological trends-described more than 2,300  years ago by Theophrastus-is the association of small leaves with dry and cold climates, which has recently been recognized for eudicotyledonous plants at a global scale1-3. For eudicotyledons, this pattern has been attributed to the fact that small leaves have a thinner boundary layer that helps to avoid extreme leaf temperatures4 and their leaf development results in vein traits that improve water transport under cold or dry climates5,6. However, the global distribution of leaf size and its adaptive basis have not been tested in the grasses, which represent a diverse lineage that is distinct in leaf morphology and that contributes 33% of terrestrial primary productivity (including the bulk of crop production)7. Here we demonstrate that grasses have shorter and narrower leaves under colder and drier climates worldwide. We show that small grass leaves have thermal advantages and vein development that contrast with those of eudicotyledons, but that also explain the abundance of small leaves in cold and dry climates. The worldwide distribution of leaf size in grasses exemplifies how biophysical and developmental processes result in convergence across major lineages in adaptation to climate globally, and highlights the importance of leaf size and venation architecture for grass performance in past, present and future ecosystems.