We have analyzed expression conferred by five subdomains of the cauliflower mosaic virus (CaMV) 35S enhancer in mature transgenic plants. Expression was detected from subdomains that gave no expression at earlier stages of development indicating developmental regulation of expression and confirming the modular organization of the enhancer. In several cases the expression patterns are highly restricted in cell type, providing useful markers for developmental studies. Comparison of expression patterns conferred by various combinations of 35S enhancer cis-elements suggests that synergistic interactions among cis-elements may play an important role in defining tissue-specific expression. This has implications for the nature of a cis-element combinatorial code that could define expression throughout development.