Cytokinin regulates many important aspects of plant development in aerial and subterranean organs. The hormone is part of an intrinsic genetic network controlling organ development and growth in these two distinct environments that plants have to cope with. Cytokinin also mediates the responses to variable extrinsic factors, such as light conditions in the shoot and availability of nutrients and water in the root, and has a role in the response to biotic and abiotic stress. Together, these activities contribute to the fine-tuning of quantitative growth regulation in plants. We review recent progress in understanding the cytokinin system and its links to the regulatory pathways that respond to internal and external signals.