Bowen, A. E.; Hadjivasiliou, Z.

Developmental patterns can scale with size during growth, a phenomenon commonly attributed to morphogen scaling. Although patterning is orchestrated by gene regulatory networks (GRNs) activated by morphogens, how GRN dynamics interact with growth is not understood. We present a theoretical framework that integrates morphogen signalling, GRN dynamics, and tissue growth. We show that pattern scaling emerges from the interplay of GRN dynamics and growth, even in the absence of morphogen scaling. This relies on memory effects encoded in the GRNs, providing a cell- autonomous route to global scaling, and offering a general mechanism for size-invariant patterning beyond morphogen-based models.