Whole-cell lysates from cells in monolayer culture and 3D organoids were prepared as described previously (17 (link)). Equivalent amounts (20–40 μg) of protein were loaded into a NuPAGE 4 to 12% Bis-Tris gel. Following electrophoresis, transfer to a polyvinylidene difluoride membrane, and blocking with 5% bovine serum albumin or non-fat milk, membranes were incubated with primary antibodies at 4°C overnight. The primary antibodies used were follows: anti-LOX (1:500; NB100–2527; Novus Biologicals, Centennial, CO, USA), anti-TP63 (1:1000; ab124762; Abcam, Cambridge, UK), anti-IVL (1:1000; I9018; Millipore Sigma, Burlington, MA, USA), anti-DSG1 (1:1000; sc-137164; Santa Cruz Biotechnology, Dallas, TX, USA), anti-BMP2 (1:1000; ab214821; Abcam), anti-phospho-SMAD1/5/9 (1:1000;13820S; Cell Signaling Technology, Danvers, MA, USA), and anti-β-actin (1:5000; A5316; Millipore Sigma). Immunoblots were detected with an appropriate horseradish peroxidase (HRP)-conjugated secondary antibody (1:2000; NA934 or NA 931; Amersham BioSciences, Buckinghamshire, UK) by ECL detection (Bio-Rad Laboratories, Hercules, CA, USA). β-Actin served as a loading control.