I used the theory of generalized gradient flows to answer this question. The resulting theoretical framework (cellular gradient flow) could explain single-cell-level rules (e.g., unidirectional differentiation) in multicellular systems as a result of population-level goal-directed behavior.
Shuhei A. Horiguchi, Tetsuya J. Kobayashi. Cellular gradient flow structure linking single-cell-level rules and population-level dynamics, Phys. Rev. Research 5, L022052, (2023)