Proposed model of GA metabolism during barley grain germination. The uptake of water into the grain initiates active GA metabolism in the scutellar epithelial layer, where stored precursors ent-kaurene (KAU) and GA53 are converted to GA1, which is subsequently glycosylated to form GA1-G. The GA1-G, KAU, and GA53 are secreted and diffuse to the aleurone layer, where KAU and GA53 again act as precursor metabolites for active GA synthesis. The GA1-G is de-glycosylated and free GA1 causes transcription of GA3ox, GA13ox, and GA20x genes. The GA3 so produced may be glycosylated to form GA3-G before it is secreted, although it is also possible that free GA3 is secreted. The GA3-G, GA3, and GA4 from the aleurone layer diffuse to adjacent aleurone cells, where KAU and GA53, and some GA1-G have also diffused from the scutellum. The process continues from the proximal end towards the distal end of the aleurone layer; the concentration of the GA1-G is likely to become limiting towards the distal end of the grain, while GA3-G, GA3, and GA4 will be replenished en route. The long-distance transport of the hydrophobic ent-kaurene may involve lipid transport proteins. The red stars indicate gene transcription.