Cellular Potts Model — Surface Tension Cell Sorting
Steinberg's differential adhesion hypothesis: cells sort by adhesion energy
Step: 0
Energy: —
High J(A-B) vs J(A-A),J(B-B) → cell sorting (DAH)
■ Type A ■ Type B ■ Medium
Differential Adhesion Hypothesis (Steinberg 1963): Cell sorting emerges purely from thermodynamic minimization of surface energy. Cells with stronger homotypic adhesion engulf cells with weaker. Hamiltonian: H = Σ J(σᵢ,σⱼ)(1−δ) + κ Σₛ (aₛ−A₀)² — interface energy + area conservation. Metropolis rule: copy spin with prob min(1, exp(−ΔH/T)). Higher T → more fluctuations, less sorting.
Key prediction: if J(A-B) > J(A-A), J(B-B) → A surrounds B (lower overall surface energy). Confirmed in zebrafish germ layers.