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In the stems of terrestrial vascular plants studied to date, the diameter of xylem water-conducting conduits D widens predictably with distance from the stem tip L approximating D ∝ Lb , with b ≈ 0.2. Because conduit diameter is central for conductance, it is essential to understand the cause of this remarkably pervasive pattern. We give reason to suspect that tip-to-base conduit widening is an adaptation, favored by natural selection because widening helps minimize the increase in hydraulic resistance that would otherwise occur as an individual stem grows longer and conductive path length increases. Evidence consistent with adaptation includes optimality models that predict the 0.2 exponent. The fact that this prediction can be made with a simple model of a single capillary, omitting much biological detail, itself makes numerous important predictions, e.g. that pit resistance must scale isometrically with conduit resistance. The idea that tip-to-base conduit widening has a nonadaptive cause, with temperature, drought, or turgor limiting the conduit diameters that plants are able to produce, is less consistent with the data than an adaptive explanation. We identify empirical priorities for testing the cause of tip-to-base conduit widening and underscore the need to study plant hydraulic systems leaf to root as integrated wholes.

Temperature is the most important factor affecting growth at high altitudes. As trees use much of the allocated carbon gained from photosynthesis to produce branches and stems, information on the timing and dynamics of secondary wood growth is crucial to assessing temperature thresholds for xylogenesis. We have carried out histological analyses to determine cambial activity and xylem cell differentiation in conifers growing at the treeline on the eastern Alps in two sites during 2002- 2004 with the aim of linking the growth process with temperature and, consequently, of defining thresholds for xylogenesis. Cambial activity occurred from May to July-August and cell differentiation from May-June to September-October. The earliest start of radial enlargement was observed in stone pine in mid-May, while Norway spruce was the last species to begin tracheid differentiation. The duration of wood formation varied from 90 to 137 days, depending on year and site, with no difference between species. Longer durations were observed in trees on the south-facing site because of the earlier onset and later ending of cell production and differentiation. The threshold temperatures at which xylogenesis had a 0.5 probability of being active were calculated by logistic regressions. Xylogenesis was active when the mean daily air temperature was 5.6-8.5°C and mean stem temperature was 7.2-9°C. The similar thresholds among all trees suggested the existence of thermal limits in wood formation that correspond with temperatures of 6-8°C that are supposed to limit growth at the treeline. Different soil temperature thresholds between sites indicated that soil temperature may not be the main factor limiting xylogenesis. This study represents the first attempt to define a threshold through comparative assessment of xylem growth and tissue temperatures in stem meristems at high altitudes.

Thermal dissipation probe (TDP) method (Granier, 1985) is widely used to estimate tree transpiration (i.e., the water evaporated from the leaves) because it is simple to build, easy to install, and relatively inexpensive. However, the universality of the original calibration has been questioned and, in many cases, proved to be inaccurate. Thus, when the TDP is used in a new species, specific tests should be carried out. Our aim was to propose a new method for improving the accuracy of TDP on trees in the field. Small hazelnut trees (diameter at breast height 5 cm) were used for the experiment. The response of TDP sensors was compared with a reference water uptake measured with an electronic potometer system provided with a high precision liquid flow meter. We equipped three stems where we measured the sap flow density, the sapwood area (by using fuchsine), the total tree water uptake (reference), and the main meteorological parameters during summer 2018. Results confirmed that the original Granier’s calibration underestimated the effective tree transpiration (relative error about −60%). We proposed a new equation for improving the measurement accuracy within an error of about 4%. The system proposed appeared an easier solution compared to potted trees and particularly suitable for orchards, thus contributing to improve the irrigation management worldwide.

Leaves of vascular plants use specific tissues to irrigate the lamina (veins) and to regulate water loss (stomata), to approach homeostasis in leaf hydration during photosynthesis. As both tissues come with attendant costs, it would be expected that the synthesis and spacing of leaf veins and stomata should be coordinated in a way that maximizes benefit to the plant. We propose an innovative geoprocessing method based on image editing and a geographic information system to study the quantitative relationships between vein and stomatal spatial patterns on leaves collected from 31 angiosperm species from different biomes. The number of stomata within each areole was linearly related to the length of the looping vein contour. As a consequence of the presence of free‐ending veinlets, the minimum mean distance of stomata from the nearest veins was invariant with areole size in most of the species, and species with smaller distances carried a higher density of stomata. Uniformity of spatial patterning was consistent within leaves and species. Our results demonstrate the existence of an optimal spatial organization of veins and stomata, and suggest their interplay as a key feature for achieving a constant mesophyll hydraulic resistance throughout the leaf.

Global climate change-induced droughts are provoking events of forest mortality worldwide, with loss of tree biomass and consequent ecosystem services. Ameliorating the effects of drought requires understanding the causes of forest mortality, with failure of the hydraulic system being an important contributor. Comparative anatomical data strongly suggest that, all else being equal, wider conduits are more vulnerable to drought-induced embolism than narrow ones. However, physiology experiments do not provide consistent support for such a link. If a vulnerability-diameter link exists, though, it would contribute not only to explaining and predicting forest mortality but also to interventions to render individual trees more drought resistant. Given that xylem conduits scale with plant height, taller plants have wider conduits. If there is a vulnerability-diameter link, then this would help explain why taller plants are often more vulnerable to climate change-induced drought. Links between conduit diameter, plant height, and vulnerability would also provide guidance for standardizing sampling of hydraulic variables across individuals and suggest that selecting for relatively narrow conduits at given height from the tree top could produce more drought resistant varieties. As a result, given current ambiguities, together with the potential importance of a link, it is important to maintain the vulnerability-diameter link as a research priority.

The radial growth of plant stem is based on the development of cribro-vascular cambium tissues. It affects the transport efficiency of water, mineral nutrients and photoassimilates and, ultimately, also plant height. The rate of cambial cell divisions for the assembly of new xylem and phloem tissue primordia and the rate of differentiation of the primordia into mature tissues determine the amount of biomass produced and, in the case of woody species, the wood quality. These complex physiological processes proceed at a rate which depends on several factors, acting at various levels: growth regulators, resource availability and environmental factors. Several hormonal signals and, more recently, further regulatory molecules, have been shown to be involved in the induction and maintenance of cambium and the formation of secondary vascular tissues. The control of xylem cell patterning is of particular interest, because it determines the diameter of xylem vessels, which is central to the efficiency of water and nutrient transport from roots to leaves through the stem and may strongly influence the growth in height of the tree. Increasing scientific evidence have proved the role of other hormones in cambial cell activities and the study of the hormonal signals and their crosstalking in cambial cells may foster our understanding of the dynamics of xylogenesis and of the mechanism of vessel size control along the stem. In this article, the role of the hormonal signals involved in the control of cambium and xylem development in trees and their crosstalking are reviewed.

Bark thickness is ecologically crucial, affecting functions from fire protection to photosynthesis. Bark thickness scales predictably with stem diameter, but there is little consensus on whether this scaling is a passive consequence of growth or an important adaptive phenomenon requiring explanation. With a comparative study across 913 species, we test the expectation that, if bark thickness–stem diameter scaling is adaptive, it should be possible to find ecological situations in which scaling is predictably altered, in this case between species with different types and deployments of phloem. ‘Dicots’ with successive cambia and monocots, which have phloem-free bark, had predictably thinner inner (mostly living) bark than plants with single cambia. Lianas, which supply large leaf areas with limited stem area, had much thicker inner bark than self-supporting plants. Gymnosperms had thicker outer bark than angiosperms. Inner bark probably scales with plant metabolic demands, for example with leaf area. Outer bark scales with stem diameter less predictably, probably reflecting diverse adaptive factors; for example, it tends to be thicker in fire-prone species and very thin when bark photosynthesis is favored. Predictable bark thickness–stem diameter scaling across plants with different photosynthate translocation demands and modes strongly supports the idea that this relationship is functionally important and adaptively significant.

A remarkable increase in vapor pressure deficit (VPD) has been recorded in the last decades in relation to global warming. Higher VPD generally leads to stomatal closure and limitations to leaf carbon uptake. Assessing tree conductance responses to VPD is a key step for modeling plant performances and productivity under future environmental conditions, especially when trees are cultivated well outside their native range as for hazelnut ( Corylus spp.). Our main aim is to assess the stand-level surface canopy conductance ( G surf ) responses to VPD in hazelnut across different continents to provide a proxy for potential productivity. Tree sap flow ( Fd ) was measured by Thermal dissipation probes (TDP) probes (six per sites) in eight hazelnut orchards in France, Italy, Georgia, Australia, and Chile during three growing seasons since 2016, together with the main meteorological parameters. We extracted diurnal Fd to estimate the canopy conductance G surf. . In all the sites, the maximum G surf occurred at low values of VPD (on average 0.57 kPa) showing that hazelnut promptly avoids leaf dehydration and that maximum leaf gas exchange is limited at relatively low VPD (i.e., often less than 1 kPa). The sensitivity of the conductance vs . VPD (i.e., -d G /dlnVPD) resulted much lower (average m = −0.36) compared to other tree species, with little differences among sites. We identified a range of suboptimal VPD conditions for G surf maximization ( G surf > 80% compared to maximum) in each site, named “VPD 80 ,” which multiplied by the mean G surf might be used as a proxy for assessing the maximum gas exchange of the orchard with a specific management and site. Potential gas exchange appeared relatively constant in most of the sites except in France (much higher) and in the driest Australian site (much lower). This study assessed the sensitivity of hazelnut to VPD and proposed a simple proxy for predicting the potential gas exchange in different areas. Our results can be used for defining suitability maps based on average VPD conditions, thus facilitating correct identification of the potentially most productive sites.

The diameter of vascular conduits increases towards the stem base. It has been suggested that this profile is an efficient anatomical feature for reducing the hydraulic resistance when trees grow taller. However, the mechanism that controls the cell diameter along the plant is not fully understood. The timing of cell differentiation along the stem was investigated. Cambial activity and cell differentiation were investigated in aPicea abiestree (11.5 m in height) collecting microsamples at nine different heights (from 1 to 9 m) along the stem with a 4 d time interval. Wood sections (8–12 μm thick) were stained and observed under a light microscope with polarized light to differentiate the developing xylem cells. Cell wall lignification was detected using cresyl violet acetate. The first enlarging cells appeared almost simultaneously along the tree axis indicating that cambium activation is not heightdependent. A significant increase in the duration of the cell expansion phase was observed towards the tree base: at 9 m from the ground, xylem cells expanded for 7 d, at 6 m for 14 d, and at 3 m for 19 d. The duration of the expansion phase is positively correlated with the lumen area of the tracheids (r² =0.68,P< 0.01) at the same height. By contrast, thickness of the cell wall of the earlywood did not show any trend with height. The lumen area of the conduits down the stem appeared linearly dependent on time during which differentiating cells remained in the expansion phase. However, the inductive signal of such long-distance patterned differentiation remains to be identified.

• 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.