Angiosperm hydraulic performance is crucially affected by the diameters of vessels, the water conducting conduits in the wood. Hydraulic optimality models suggest that vessels should widen predictably from stem tip to base, buffering hydrodynamic resistance accruing as stems, and therefore conductive path, increase in length. Data from 257 species (609 samples) show that vessels widen as predicted with distance from the stem apex across angiosperm orders, habits and habitats. Standardising for stem length, vessels are only slightly wider in warm/moist climates and in lianas, showing that, rather than climate or habit, plant size is by far the main driver of global variation in mean vessel diameter. Terminal twig vessels become wider as plant height increases, while vessel density decreases slightly less than expected tip to base. These patterns lead to testable predictions regarding evolutionary strategies allowing plants to minimise carbon costs per unit leaf area even as height increases.
Within a tree the lumen of the xylem conduits varies widely (by at least 1 order of magnitude). Transversally in the stem conduits are smaller close to the pith and larger in the outermost rings. Axially (i.e. from petioles to roots) conduits widen from the stem apex downwards in the same tree ring. This axial variation is proposed as being the most efficient anatomical adjustment for stabilizing hydraulic path-length resistance with the progressive growth in height. The hydrodynamic (i.e. physical) constraint shapes the whole xylem conduits column in a very similar way in different species and environments. Our aim is to provide experimental evidence that the axial conduit widening is an ineluctable feature of the vascular system in plants. If evolution has favoured efficient distribution networks (i.e. total resistance is tree-size independent) the axial conduit widening can be predicted downwards along the stem. Indeed, in order to compensate for the increase in path length with growth in height the conduit size should scale as a power function of tree height with an exponent higher than 0.2. Similarly, this approach could be applied in branches and roots but due to the different lengths of the path roots-leaves the patterns of axial variations of conduit size might slightly deviate from the general widening trend. Finally, we emphasize the importance of sampling standardization with respect to tree height for cor- rectly comparing the anatomical characteristics of different individuals
Early observations led Sanio [Wissen. Bot., 8, (1872) 401] to state that xylem conduit diameters and lengths in a coniferous tree increase from the apex down to a height below which they begin to decrease towards the tree base. Sanio's law of vertical tapering has been repeatedly tested with contradictory results and the debate over the scaling of conduit diameters with distance from the apex has not been settled. The debate has recently acquired new vigour, as an accurate knowledge of the vertical changes in wood anatomy has been shown to be crucial to scaling metabolic properties to plant and ecosystem levels. Contrary to Sanio's hypothesis, a well known model (MST, metabolic scaling theory) assumes that xylem conduits monotonically increase in diameter with distance from the apex following a power law. This has been proposed to explain the three-fourth power scaling between size and metabolism seen across plants. Here, we (i) summarized available data on conduit tapering in trees and (ii) propose a new numerical model that could explain the observed patterns. Data from 101 datasets grouped into 48 independent profiles supported the notions that phylogenetic group (angiosperms versus gymnosperms) and tree size strongly affected the vertical tapering of conduit diameter. For both angiosperms and gymnosperms, within-tree tapering also varied with distance from the apex. The model (based on the concept that optimal conduit tapering occurs when the difference between photosynthetic gains and wall construction costs is maximal) successfully predicted all three major empirical patterns. Our results are consistent with Sanio's law only for large trees and reject the MST assumptions that vertical tapering in conduit diameter is universal and independent of rank number.
Summary• Vertical conduit tapering is proposed as an effective mechanism to almost eliminate the increase in hydraulic resistance with increased height. Despite this potential role, very little is known about its changes during ontogeny.• Here, conduit tapering and stem morphology of young/small and old/tall individuals of Acer pseudoplatanus in the field, as well as 3-yr-old grafted trees from both age classes, were analysed. The distribution of hydraulic resistance along stems was also determined in a subsample of trees.• Substantial conduit tapering was found in small trees (field-grown and grafted from both age classes), whereas values were lower in tall trees, indicating that tapering was a size-related, not an age-related process. Apical conduit diameters were larger in tall trees and were inversely correlated with the degree of tapering. Hydraulic resistance increased less than linearly with distance from the apex. Its scaling against distance was indistinguishable from that predicted from anatomical measurements.• Conduit tapering was an effective but partial mechanism of compensation for the increase in hydraulic resistance with tree height. Size-related changes in tapering and in apical conduit diameters may be explained by the combined need to reduce the buildup of hydraulic resistance while minimizing the carbon costs of building vessel walls.
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