Multi-scale phenotyping of leaf expansion in response to environmental changes: the whole is more than the sum of parts.

The leaf is a multi-scale dynamic unit that is determined by mechanisms at different organizational scales (cell, tissue, whole leaf and whole plant) and affected by both internal (genotype) and external (environmental) determinisms. The recent development of phenotyping platforms and imaging techniques provides new insights into the temporal and spatial patterns of leaf growth as affected by those determinisms. Conclusions about the overriding mechanisms often depend on the considered organizational scale and of time resolution which varies from minutes to several weeks. Analyses of leaf growth responses to environmental conditions have revealed robust emerging properties at whole plant or whole leaf scales. They have highlighted that the control of individual leaf expansion is more complex than merely the sum of cellular processes, and the control at the whole plant level is more complex than the sum of individual leaf expansions. However, in many cases, the integrated leaf-growth variable can be simplified to a limited set of underlying variables to be measured for comparative analyses of leaf growth or modelling purposes.