Several plastid signals have been identified that regulate the transcription of nuclear genes for plastid and non-plastid proteins. These signals are related to the photosynthetic metabolism of chloroplasts and include porphyrins, reactive oxygen intermediates and carotenoids. The metabolic and developmental state of the chloroplast also control cell differentiation and leaf morphogenesis, but the signaling pathways have not been characterized. Plastid-to-nucleus and light-signaling pathways are separable in some but not all cases. Retrograde signaling thus plays a central role in coordinating gene expression in the nucleus, plastid and mitochondrion, and in integrating pathways of cellular metabolism and development.