Summary1. The impacts of elevated atmospheric CO 2 and/or O 3 have been examined over 4 years using an open-air exposure system in an aggrading northern temperate forest containing two different functional groups (the indeterminate, pioneer, O 3 -sensitive species Trembling Aspen, Populus tremuloides and Paper Birch, Betula papyrifera , and the determinate, late successional, O 3 -tolerant species Sugar Maple, Acer saccharum ). 2. The responses to these interacting greenhouse gases have been remarkably consistent in pure Aspen stands and in mixed Aspen/Birch and Aspen/Maple stands, from leaf to ecosystem level, for O 3 -tolerant as well as O 3 -sensitive genotypes and across various trophic levels. These two gases act in opposing ways, and even at low concentrations (1·5 × ambient, with ambient averaging 34 -36 nL L − 1 during the summer daylight hours), O 3 offsets or moderates the responses induced by elevated CO 2 . 3. After 3 years of exposure to 560 µ mol mol − 1 CO 2 , the above-ground volume of Aspen stands was 40% above those grown at ambient CO 2 , and there was no indication of a diminishing growth trend. In contrast, O 3 at 1·5 × ambient completely offset the growth enhancement by CO 2 , both for O 3 -sensitive and O 3 -tolerant clones. Implications of this finding for carbon sequestration, plantations to reduce excess CO 2 , and global models of forest productivity and climate change are presented.
Some anatomical characteristics in leaves relating to hydraulic conductance and stomatal conductance were examined in six temperate deciduous tree species. The fourth power of the radius of the conducting elements in xylem (r 4 ) and the area of mesophyll and epidermal cells per unit length of leaf cross-section (u) were high in leaves with high hydraulic conductance (L). Stomatal conductance (g s ) and stomatal sensitivity to an increase in leaf water potential (s i ) correlated positively with the length of stomatal pore (l), but negatively with the guard cell width (z) and the length of the dorsal side of the guard cells (l d ). Stomatal sensitivity to a decrease in leaf water potential (s d ) correlated negatively with l and positively with z and l d. The anatomical characteristics associated with hydraulic conductance (r 4 and u) and those associated with stomatal conductance and sensitivity to changes of leaf water potential (l, z and l d ) were correlated. We conclude that hydraulic conductance may depend on anatomical characteristics of xylem, mesophyll and epidermis, and stomatal conductance and its sensitivity to changing water potential may depend on anatomical characteristics of stomata. The correlation of shoot hydraulic conductance with stomatal conductance and its sensitivity may be based largely on the correlation between the anatomical characteristics of the water conducting system and stomata in these trees.
Leaf gas exchange parameters and the content of ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Rubisco) in the leaves of two 2‐year‐old aspen (Populus tremuloides Michx.) clones (no. 216, ozone tolerant and no. 259, ozone sensitive) were determined to estimate the relative stomatal and mesophyll limitations to photosynthesis and to determine how these limitations were altered by exposure to elevated CO2 and/or O3. The plants were exposed either to ambient air (control), elevated CO2 (560 p.p.m.) elevated O3 (55 p.p.b.) or a mixture of elevated CO2 and O3 in a free air CO2 enrichment (FACE) facility located near Rhinelander, Wisconsin, USA. Light‐saturated photosynthesis and stomatal conductance were measured in all leaves of the current terminal and of two lateral branches (one from the upper and one from the lower canopy) to detect possible age‐related variation in relative stomatal limitation (leaf age is described as a function of leaf plastochron index). Photosynthesis was increased by elevated CO2 and decreased by O3 at both control and elevated CO2. The relative stomatal limitation to photosynthesis (ls) was in both clones about 10% under control and elevated O3. Exposure to elevated CO2 + O3 in both clones and to elevated CO2 in clone 259, decreased ls even further – to about 5%. The corresponding changes in Rubisco content and the stability of Ci/Ca ratio suggest that the changes in photosynthesis in response to elevated CO2 and O3 were primarily triggered by altered mesophyll processes in the two aspen clones of contrasting O3 tolerance. The changes in stomatal conductance seem to be a secondary response, maintaining stable Ci under the given treatment, that indicates close coupling between stomatal and mesophyll processes.
At northern latitudes a rise in atmospheric humidity and precipitation is predicted as a consequence of global climate change. We studied several growth and functional traits of hybrid aspen (Populus tremula L.×P. tremuloides Michx.) in response to elevated atmospheric humidity (on average 7% over the ambient level) in a free air experimental facility during three growing seasons (2008–2010) in Estonia, which represents northern temperate climate (boreo-nemoral zone). Data were collected from three humidified (H) and three control (C) plots, and analysed using nested linear models. Elevated air humidity significantly reduced height, stem diameter and stem volume increments and transpiration of the trees whereas these effects remained highly significant also after considering the side effects from soil-related confounders within the 2.7 ha study area. Tree leaves were smaller, lighter and had lower leaf mass per area (LMA) in H plots. The magnitude and significance of the humidity treatment effect – inhibition of above-ground growth rate – was more pronounced in larger trees. The lower growth rate in the humidified plots can be partly explained by a decrease in transpiration-driven mass flow of NO3
− in soil, resulting in a significant reduction in the measured uptake of N to foliage in the H plots. The results suggest that the potential growth improvement of fast-growing trees like aspens, due to increasing temperature and atmospheric CO2 concentration, might be smaller than expected at high latitudes if a rise in atmospheric humidity simultaneously takes place.
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