Trees are among the best natural archives of past environmental information. Xylem anatomy preserves information related to tree allometry and ecophysiological performance, which is not available from the more customary ring-width or wood-density proxy parameters. Recent technological advances make tree-ring anatomy very attractive because time frames of many centuries can now be covered. This calls for the proper treatment of time series of xylem anatomical attributes. In this article, we synthesize current knowledge on the biophysical and physiological mechanisms influencing the short- to long-term variation in the most widely used wood-anatomical feature, namely conduit size. We also clarify the strong mechanistic link between conduit-lumen size, tree hydraulic architecture and height growth. Among the key consequences of these biophysical constraints is the pervasive, increasing trend of conduit size during ontogeny. Such knowledge is required to process time series of anatomical parameters correctly in order to obtain the information of interest. An appropriate standardization procedure is fundamental when analysing long tree-ring-related chronologies. When dealing with wood-anatomical parameters, this is even more critical. Only an interdisciplinary approach involving ecophysiology, wood anatomy and dendrochronology will help to distill the valuable information about tree height growth and past environmental variability correctly.

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 correctly comparing the anatomical characteristics of different individuals.

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 a Picea abies tree (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 height-dependent. 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.