Mechanics of cell specialization. (A1, A2, and B) The relationship between spatial (form) and transcriptional (function) networks over time during cell specialization. Alignment of the networks, where the architecture and transcriptional networks become mutually related. Prior to the aligned state, we propose two possible models: (A1) overall coregulated gene content emerges according to overall chromosome associations, or form precedes function, and (A2) the transcriptional network and coregulated genes precede and shape the architecture network, or form follows function. (B) After alignment, feedback between form and function networks allows fine-tuning to achieve optimal cell-specific network configurations.