“…The testa gradually becomes impermeable during the later stages of seed maturation (Gillikin and Graham, 1991); the biochemical basis of this hardening varies among families and species. Testa thickness is influential (Miao et al, 2001), but is not the sole basis of impermeability (de Souza and Marcos-Filho, 2001), which is also affected by the testa's composition (Smýkal et al, 2014;Cechová et al, 2017;Hradilová et al, 2017). Cell wall thickness in the outer integuments varies throughout seed development (Verdier et al, 2013), along with the accumulation of polyphenolic compounds; oxidation products of the latter are thought to contribute to impermeability in both barrel medic (Medicago truncatula) and pea (Pisum sativum) (Marbach and Mayer, 1974;Werker et al, 1979;Moïse et al, 2005), but also modifies permeability in Arabidopsis (Debeaujon et al, 2000).…”