Mechanisms that coordinate growth during development are essential for producing animals with proper organ proportion. Here we describe a pathway through which tissues communicate to coordinate growth. During Drosophila melanogaster larval development, damage to imaginal discs activates a regeneration checkpoint through expression of Dilp8. This both produces a delay in developmental timing and slows the growth of undamaged tissues, coordinating regeneration of the damaged tissue with developmental progression and overall growth. Here we demonstrate that Dilp8-dependent growth coordination between regenerating and undamaged tissues, but not developmental delay, requires the activity of nitric oxide synthase (NOS) in the prothoracic gland. NOS limits the growth of undamaged tissues by reducing ecdysone biosynthesis, a requirement for imaginal disc growth during both the regenerative checkpoint and normal development. Therefore, NOS activity in the prothoracic gland coordinates tissue growth through regulation of endocrine signals.KEYWORDS regeneration; nitric oxide; ecdysone; allometry; growth control A LLOMETRY, broadly defined as the scaling of organ growth, can have a profound impact on the biological function in animals. For example, in the male dung beetle, Onthophagus netriventris, an inverse allometry is observed between horn and testes size, producing distinct reproductive strategies (Simmons and Emlen 2006;Emlen et al. 2012). Allometric growth regulation can also impact human health where variation from optimal relative heart size can increase susceptibility to cardiovascular disease (Hill and Olson 2008). Despite the fundamental role of growth scaling in biology, no described pathways explain how tissues coordinate growth. Our understanding of growth regulation has been focused on either tissue-autonomous mechanismssuch as how morphogens regulate the activity of cellular growth pathways-or systemic mechanisms such as how endocrine factors control growth in response to environmental change.These tissue-autonomous and systemic pathways of growth regulation do not explain allometric growth observed during development. Transplantation experiments (Madhavan and Schneiderman 1969) and growth perturbation experiments in Drosophila and other insects (Nijhout and Emlen 1998;Simmons and Emlen 2006;Parker and Shingleton 2011) suggest that interorgan communication may be necessary for allometric growth. Based on these observations, Stern and Emlen (1999) proposed a model for growth coordination that requires communication between growing organs, either directly or indirectly through an endocrine organ. However, the mechanism of this communication pathway has been unclear.In Drosophila larvae, the growth of the imaginal discs is tightly regulated to produce adult structures with specific size and proportion (Mirth and Shingleton 2012;Callier and Nijhout 2013). Allometry between these tissues is preserved even when developmental growth programs are altered. For example, Drosophila imaginal discs have a remar...