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 degrees C and mean stem temperature was 7.2-9 degrees C. The similar thresholds among all trees suggested the existence of thermal limits in wood formation that correspond with temperatures of 6-8 degrees 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.

Summary• A recent theoretical model (the West, Brown and Enquist, WBE model) hypothesized that plants have evolved a network of xylem conduits with a tapered structure (narrower conduits distally) which should minimize the cost of water transport from roots to leaves. Specific measurements are required to test the model predictions.• We sampled both angiosperms and gymnosperms (50 trees) growing in different environments with heights ranging from 0.5 to 44.4 m, measuring variations of the xylem-conduit diameter from tree top to stem base.• In all trees measured, mean hydraulically weighted conduit diameters ( Dh ) at the tree top were narrower than those at the stem base. In actively growing trees, the longitudinal variation of Dh showed a degree of tapering in agreement with WBE predictions, while trees close to their maximum height showed slightly lower conduit tapering. Comparing different species, a very good correlation was observed between degree of xylem tapering and tree height ( r 2 = 0.88; P < 0.0001) independently of any other variable (age, site, altitude, etc.).• As predicted by WBE, sampled trees seemed to converge towards similar xylem conduit tapering. However, trees approaching their maximum height had a nonoptimal tapering which appeared insufficient to compensate for the progressive increase in tree height.

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Summary• 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 highaltitude 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.

Trees play a key role in the global hydrological cycle and measurements performed with the thermal dissipation method (TDM) have been crucial in providing whole-tree water-use estimates. Yet, different data processing to calculate whole-tree water use encapsulates uncertainties that have not been systematically assessed. We quantified uncertainties in conifer sap flux density (F ) and stand water use caused by commonly applied methods for deriving zero-flow conditions, dampening and sensor calibration. Their contribution has been assessed using a stem segment calibration experiment and 4 yr of TDM measurements in Picea abies and Larix decidua growing in contrasting environments. Uncertainties were then projected on TDM data from different conifers across the northern hemisphere. Commonly applied methods mostly underestimated absolute F . Lacking a site- and species-specific calibrations reduced our stand water-use measurements by 37% and induced uncertainty in northern hemisphere F . Additionally, although the interdaily variability was maintained, disregarding dampening and/or applying zero-flow conditions that ignored night-time water use reduced the correlation between environment and F . The presented ensemble of calibration curves and proposed dampening correction, together with the systematic quantification of data-processing uncertainties, provide crucial steps in improving whole-tree water-use estimates across spatial and temporal scales.