Summary• This study aims to link three-dimensional coarse root architecture to tree stability in mature timber trees with an average of 1-m rooting depth.• Undamaged and uprooted trees were sampled in a stand damaged by a storm. Root architecture was measured by three-dimensional (3-D) digitizing. The distribution of root volume by root type and in wind-oriented sectors was analysed.• Mature Pinus pinaster root systems were organized in a rigid 'cage' composed of a taproot, the zone of rapid taper of horizontal surface roots and numerous sinkers and deep roots, imprisoning a large mass of soil and guyed by long horizontal surface roots. Key compartments for stability exhibited strong selective leeward or windward reinforcement. Uprooted trees showed a lower cage volume, a larger proportion of oblique and intermediate depth horizontal roots and less windoriented root reinforcement.• Pinus pinaster stability on moderately deep soils is optimized through a typical rooting pattern and a considerable structural adaptation to the prevailing wind and soil profile.
Intra-annual radial growth variations of two Norway spruce trees (Picea abies (L.) Karst.) were monitored over 4 years, at four heights up the stem, by means of point-dendrometers. The trees were then felled and radial wood samples were cut from the radii that had been monitored by the dendrometers and analyzed for density. From the radial growth measurements recorded by the dendrometers, we related positions within the rings to dates, thus making possible investigation of the relationships between changes within the rings in wood density and fluctuations in climate or growth rate. Radial growth started in early April and ended, with large intra-annual differences, in August or September. Short-term variations in growth rate were related to fluctuations in climate parameters and soil water reserves. The sensitivity of radial growth to climate decreased with stem height. Wood density responded strongly to drought events, and a dry period in June 1996 induced false-ring formation. Wood density was relatively independent of growth rate and climatic conditions during the first part of the growing season, but increased with decreasing radial growth rate later in the growing season.
Global biodiversity is affected by numerous environmental drivers. Yet, the extent to which global environmental changes contribute to changes in local diversity is poorly understood. We investigated biodiversity changes in a meta-analysis of 39 resurvey studies in European temperate forests (3988 vegetation records in total, 17-75 years between the two surveys) by assessing the importance of (i) coarse-resolution (i.e., among sites) vs. fine-resolution (i.e., within sites) environmental differences and (ii) changing environmental conditions between surveys. Our results clarify the mechanisms underlying the direction and magnitude of local-scale biodiversity changes. While not detecting any net local diversity loss, we observed considerable among-site variation, partly explained by temporal changes in light availability (a local driver) and density of large herbivores (a regional driver). Furthermore, strong evidence was found that presurvey levels of nitrogen deposition determined subsequent diversity changes. We conclude that models forecasting future biodiversity changes should consider coarse-resolution environmental changes, account for differences in baseline environmental conditions and for local changes in fine-resolution environmental conditions.
Variations of intrinsic water-use efficiency during the last century were investigated based on analysis of ␦ 13 C in tree rings of Abies alba from the Jura Mountains (eastern France). To separate the effects related to the age of the tree at the time the tree ring was formed from effects due to environmental changes, analyzed wood samples were extracted from a very large sample set including different tree ages and calendar dates of wood formation.For the first 75 yr of the life of Abies alba, ␦ 13 C of wood holocellulose increases with the age of the tree from Ϫ24.4‰ at age 15 to approximately Ϫ22.5‰ at age 75. Between the ages of 75 and 150 values remain constant at Ϫ22.5‰. Consequently, the effect of the tree age on isotopic discrimination has to be taken into account in studies on the long-term environmental effects on ␦ 13 C in tree rings.Divergent trends of ␦ 13 C during the last century were observed between tree rings formed at age 40 and bulk air data. The isotopic discrimination ⌬ varied insignificantly around a mean of 17.3‰ between the 1860s and the 1930s. It then decreased to 15.8‰ from the 1930s to the 1980s. Using these results and classical models of carbon discrimination, we calculated that the intrinsic water-use efficiency (A/g w , the ratio of CO 2 assimilation rate to stomatal conductance for water vapor), integrated over the year, has increased by 30% between the 1930s and the 1980s. These results, obtained at the level of mature trees, are consistent with the physiological effects of increasing CO 2 concentrations as observed in controlled experiments on young seedlings. They are consistent with the strong increases in radial growth observed for Abies alba in western Europe over the past decades. However, other long-term environmental changes such as increasing nitrogen deposition could cause similar effects.
Rising atmospheric [CO2 ], ca , is expected to affect stomatal regulation of leaf gas-exchange of woody plants, thus influencing energy fluxes as well as carbon (C), water, and nutrient cycling of forests. Researchers have proposed various strategies for stomatal regulation of leaf gas-exchange that include maintaining a constant leaf internal [CO2 ], ci , a constant drawdown in CO2 (ca - ci ), and a constant ci /ca . These strategies can result in drastically different consequences for leaf gas-exchange. The accuracy of Earth systems models depends in part on assumptions about generalizable patterns in leaf gas-exchange responses to varying ca . The concept of optimal stomatal behavior, exemplified by woody plants shifting along a continuum of these strategies, provides a unifying framework for understanding leaf gas-exchange responses to ca . To assess leaf gas-exchange regulation strategies, we analyzed patterns in ci inferred from studies reporting C stable isotope ratios (δ(13) C) or photosynthetic discrimination (∆) in woody angiosperms and gymnosperms that grew across a range of ca spanning at least 100 ppm. Our results suggest that much of the ca -induced changes in ci /ca occurred across ca spanning 200 to 400 ppm. These patterns imply that ca - ci will eventually approach a constant level at high ca because assimilation rates will reach a maximum and stomatal conductance of each species should be constrained to some minimum level. These analyses are not consistent with canalization toward any single strategy, particularly maintaining a constant ci . Rather, the results are consistent with the existence of a broadly conserved pattern of stomatal optimization in woody angiosperms and gymnosperms. This results in trees being profligate water users at low ca , when additional water loss is small for each unit of C gain, and increasingly water-conservative at high ca , when photosystems are saturated and water loss is large for each unit C gain.
Ocellulose were different between years and individuals, and mostly captured by the model, suggesting that the single-substrate hypothesis is a good approximation for tree ring studies on Pinus pinaster, at least for the environmental conditions covered by this study. A sensitivity analysis revealed that the model was mostly affected by five isotopic discrimination factors and two leaf gas-exchange parameters. Modelled early wood signals were also very sensitive to the date when cell wall thickening begins (twt). Our model could therefore be used to reconstruct twt time series and improve our understanding of how climate influences this key parameter of xylogenesis.
Static winching tests were carried out in order to determine the mechanical resistance of Maritime pine to overturning. The tested stands were selected according to podzolic soil conditions: "wet Lande", characterised by a shallow ground water table and a hard pan horizon, and "dry Lande", with a deeper ground water table and a hard pan absent or broken up. As this soil horizon limits the vertical growth of tree roots, anchorage resistance was investigated with regards to the presence or absence of a hard pan underneath each tree. To determine if mechanical behaviour differed within a stand, trees from inside the stand and edge trees at the border exposed to prevailing winds were also tested. The critical turning moment (TM crit,total ) at the base of the stem was positively related to the variable (H × DBH 2 ) (H, total tree height; DBH, tree diameter). Linear regression analyses between TM crit,total and (H × DBH 2 ) showed that the presence of a hard pan had no significant effect on anchorage resistance in uprooted trees. Stem failure occurred for 82% of trees on dry Lande when (H × DBH 2 ) < 1 m 3 . Moreover, stem failure type on dry Lande indicated that trees were better anchored. On soil with a hard pan, edge trees were found to be 20% more resistant to overturning than inner trees. Edge trees differed from inner trees in that the soil-root plate was two times larger and also possessed a larger surface area on the windward side.
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