This study demonstrates that elevated [CO 2 ] has profound effects on cell division and expansion in developing wheat (Triticum aestivum L.) leaves and on the quantitative integration of these processes in whole-leaf growth kinetics, anatomy, and carbon content. The expression of these effects, however, is modified by intrinsic factors related to genetic makeup and leaf position, and also by exposure to low vernalizing temperatures at germination. Beyond these interactions, leaf developmental responses to elevated [CO 2 ] in wheat share several remarkable features that were conserved across all leaves examined. Most significantly: (a) the contribution of [CO 2 ] effects on meristem size and activity in driving differences in whole-blade growth kinetics and final dimensions; (b) an anisotropy in cellular growth responses to elevated [CO 2 ], with final cell length and expansion in the paradermal plane being highly conserved, even when the rates and duration of cell elongation were modified, while cell cross-sectional areas were increased; (c) tissuespecific effects of elevated [CO 2 ], with significant modifications of mesophyll anatomy, including an increased extension of intercellular air spaces and the formation of, on average, one extra cell layer, while epidermal anatomy was mostly unaltered. Our results indicate complex developmental regulations of sugar effects in expanding leaves that are subjected to genetic variation and influenced by environmental cues important in the promotion of floral initiation. They also provide insights into apparently contradictory and inconsistent conclusions of published CO 2 enrichment studies in wheat.The consequences for plant growth and morphogenesis of variations in photosynthesis depend on the efficiency of the conversion of triose-P into Suc and of phloem loading at the sites of Suc production and unloading in growing sinks. Ultimately, however, they depend on the sensitivity to sugar supply of a suite of developmental processes involved in meristem initiation, cell division, expansion, and differentiation, and of the mechanisms that regulate the integration of these processes in the formation of organs with a certain shape, size, and structure. The aim of the present study was to investigate this latter area using atmospheric [CO 2 ] as a tool for manipulating sugar supply to expanding organs and wheat (Triticum aestivum L.) as a model experimental system.Based on experiments at the whole-plant or whole-leaf level, it has been reported that, relative to other species, wheat is not very responsive to elevated [CO 2 ], especially at early stages. It has been argued (Nicolas et al., 1993;Christ and Kö rner, 1995; Slafer and Rawson, 1997) that while it enhances tillering by promoting the development of axillary meristems, elevated [CO 2 ] has little to no effect on leaf development and growth in wheat, a conclusion also recently put forward for rice, another important cereal (Jitla et al., 1997). These reports challenged the evidence from other studies using wheat and a ...