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Abstract Tree growth is generally considered to be temperature limited at upper elevation treelines, yet climate factors controlling tree growth at semiarid treelines are poorly understood. We explored the influence of climate on stem growth and stable isotopes for Polylepis tarapacana Philipi, the world’s highest elevation tree species, which is found only in the South American Altiplano. We developed tree-ring width index (RWI), oxygen (δ18O) and carbon (δ13C) chronologies for the last 60 years at four P. tarapacana stands located above 4400 m in elevation, along a 500 km latitude aridity gradient. Total annual precipitation decreased from 300 to 200 mm from the northern to the southern sites. We used RWI as a proxy of wood formation (carbon sink) and isotopic tree-ring signatures as proxies of leaf-level gas exchange processes (carbon source). We found distinct climatic conditions regulating carbon sink processes along the gradient. Current growing-season temperature regulated RWI at northern-wetter sites, while prior growing-season precipitation determined RWI at arid southern sites. This suggests that the relative importance of temperature to precipitation in regulating tree growth is driven by site water availability. By contrast, warm and dry growing seasons resulted in enriched tree-ring δ13C and δ18O at all study sites, suggesting that similar climate conditions control carbon-source processes along the gradient. Site-level δ13C and δ18O chronologies were significantly and positively related at all sites, with the strongest relationships among the southern drier stands. This indicates an overall regulation of intercellular carbon dioxide via stomatal conductance for the entire P. tarapacana network, with greater stomatal control when aridity increases. This manuscript also highlights a coupling (decoupling) between physiological processes at leaf level and wood formation as a function of similarities (differences) in their climatic sensitivity. This study contributes to a better understanding and prediction of the response of high-elevation Polylepis woodlands to rapid climate changes and projected drying in the Altiplano. 

Abstract Nonstructural carbohydrates (NSCs) play a critical role in plant physiology and metabolism, yet we know little about their distribution within individual organs such as the stem. This leaves many open questions about whether reserves deep in the stem are metabolically active and available to support functional processes. To gain insight into the availability of reserves, we measured radial patterns of NSCs over the course of a year in the stemwood of temperate trees with contrasting wood anatomy (ring porous vs diffuse porous). In a subset of trees, we estimated the mean age of soluble sugars within and between different organs using the radiocarbon (14C) bomb spike approach. First, we found that NSC concentrations were the highest and most seasonally dynamic in the outermost stemwood segments for both ring-porous and diffuse-porous trees. However, while the seasonal fluctuation of NSCs was dampened in deeper stemwood segments for ring-porous trees, it remained high for diffuse-porous trees. These NSC dynamics align with differences in the proportion of functional sapwood and the arrangement of vessels between ring-porous and diffuse-porous trees. Second, radial patterns of 14C in the stemwood showed that sugars became older when moving toward the pith. The same pattern was found in the coarse roots. Finally, when taken together, our results highlight how the radial distribution and age of NSCs relate to wood anatomy and suggest that while deeper, and likely older, reserves in the stemwood fluctuated across the seasons, the deepest reserves at the center of the stem were not used to support tree metabolism under usual environmental conditions. 

Abstract Stable carbon isotope ratios (δ13C) in tree rings have been widely used to study changes in intrinsic water-use efficiency (iWUE), sometimes with limited consideration of how C-isotope discrimination is affected by tree height and canopy position. Our goals were to quantify the relationships between tree size or tree microenvironment and wood δ13C for eight functionally diverse temperate tree species in northern New England and to better understand the physical and physiological mechanisms underlying these differences. We collected short increment cores in closed-canopy stands and analyzed δ13C in the most recent 5 years of growth. We also sampled saplings in both shaded and sun-exposed environments. In closed-canopy stands, we found strong tree-size effects on δ13C, with 3.7–7.2‰ of difference explained by linear regression vs height (0.11–0.28‰ m−1), which in some cases is substantially stronger than the effect reported in previous studies. However, open-grown saplings were often isotopically more similar to large codominant trees than to shade-grown saplings, indicating that light exposure contributes more to the physiological and isotopic differences between small and large trees than does height. We found that in closed-canopy forests, δ13C correlations with diameter at breast height were nonlinear but also strong, allowing a straightforward procedure to correct tree- or stand-scale δ13C-based iWUE chronologies for changing tree size. We demonstrate how to use such data to correct and interpret multi-decadal composite isotope chronologies in both shade-regenerated and open-grown tree cohorts, and we highlight the importance of understanding site history when interpreting δ13C time series.