Mature trees have already experienced substantial increases in CO 2 concentrations during their lifetimes, and will experience continuing increases in the future. Small open-top chambers were used to enclose branchlets that were at a height of between 20 and 25 m in the canopy of the tree species Luehea seemannii Tr. & Planch. in a tropical forest in Panamá. Elevated concentrations of CO 2 increased the rate of photosynthetic carbon fixation and decreased stomatal conductance of leaves, but did not influence the growth of leaf area per chamber, the production of flower buds and fruit nor the concentration of nonstructural carbohydrates within leaves. The production of flower buds was highly correlated with the leaf area produced in the second flush of leaves, indicating that the branchlets of mature trees of Luehea seemannii are autonomous to a considerable extent. Elevated levels of CO 2 did increase the concentration of nonstructural carbohydrates in woody stem tissue. Elevated CO 2 concentration also they increased the ratio of leaf area to total biomass of branchlets, and tended to reduce individual fruit weight. These data suggest that the biomass allocation patterns of mature trees may change under future elevated levels of CO 2 . Although there were no effects on growth during the experiment, the possibility of increased growth in the season following CO 2 enrichment due to increased carbohydrate concentrations in woody tissue cannot be excluded.
Key-words: Luehea seemanniiTr. & Planch.; branch autonomy; branch-bag; carbohydrate; elevated CO 2 ; forest canopy; growth; photosynthesis.
INTRODUCTIONTrees are long-lived organisms. Tropical trees that began life in the late eighteenth century have already been exposed to considerable increases in carbon dioxide concentrations ([CO 2 ]) during their lifetimes, and will be exposed to continuing increases in the decades to come. Understanding how these mature canopy trees respond to elevated [CO 2 ], with respect to their rate of photosynthetic carbon gain, growth and reproductive output, is important because mature trees are responsible for forest carbon sequestration and provide the seed bank for forest regeneration. Few studies have attempted to investigate the effects of elevated [CO 2 ] in mature forest stands because of the difficulties involved in exposing whole trees to such concentrations and also accessing the canopy. The development of open-top chambers permitted the study of the influence of elevated [CO 2 ] in situ (Drake et al. 1989), but limited studies to vegetation types where the plants under study were similarly scaled, for example, intertidal communities (e.g. Drake et al. 1989), herbaceous communities (e.g. Potvin & Vasseur 1997), young trees (e.g. Norby, O'Neill & Luxmoore 1986;Idso, Kimball & Allen 1991;Lovelock et al. 1998), and woody communities of low stature (e.g. Cipollini, Drake & Whigham 1993). The development of free-air CO 2 enrichment (FACE) technology (Hendrey 1992) has increased the scale at which the influence of elev...