In addition to their contribution to metabolism, chloroplasts emit signals that influence the expression of nuclear genes that contribute to numerous plastidic and extraplastidic processes. Plastid-to-nucleus signalling optimizes chloroplast function, regulates growth and development, and affects responses to environmental cues. An incomplete list of plastid signals is available and particular plastid-to-nucleus signalling mechanisms are partially understood. The plastid-to-nucleus signalling that depends on the
GENOMES UNCOUPLED
(
GUN
) genes couples the expression of nuclear genes to the functional state of the chloroplast. Analyses of
gun
mutants provided insight into the mechanisms and biological functions of plastid-to-nucleus signalling.
GUN
genes contribute to chloroplast biogenesis, the circadian rhythm, stress tolerance, light signalling and development. Some have criticized the
gun
mutant screen for employing inhibitors of chloroplast biogenesis and suggested that
gun
alleles do not disrupt significant plastid-to-nucleus signalling mechanisms. Here, I briefly review
GUN
-dependent plastid-to-nucleus signalling, explain the flaws in the major criticisms of the
gun
mutant screen and review the influence of plastids on light signalling and development.