Cutinized and suberized barriers in leaves and roots: Similarities and differences

Anatomical, histochemical, chemical, and biosynthetic similarities and differences of cutinized and suberized plant cell walls are presented and reviewed in brief. Based on this, the functional properties of cutinized and suberized plant cell walls acting as transport barriers are compared and discussed in more detail. This is of general importance because fundamental misconceptions about relationships in plant-environment water relations are commonly encountered in the scientific literature. It will be shown here, that cuticles represent highly efficient apoplastic transport barriers significantly reducing the diffusion of water and dissolved compounds. The transport barrier of cuticles is mainly established by the deposition of cuticular waxes. Upon wax extraction, with the cutin polymer remaining, cuticular permeability for water and dissolved non-ionized and lipophilic solutes are increasing by 2-3 orders of magnitude, whereas polar and charged substances (e.g., nutrient ions) are only weakly affected (2- to 3-fold increases in permeability). Suberized apoplastic barriers without the deposition of wax are at least as permeable as the cutin polymer matrix without waxes and hardly offer any resistance to the free movement of water. Only upon the deposition of significant amounts of wax, as it is the case with suberized periderms exposed to the atmosphere, an efficient transport barrier for water can be established by suberized cell walls. Comparing the driving forces (gradients between water potentials inside leaves and roots and the surrounding environment) for water loss acting on leaves and roots, it is shown that leaves must have a genetically pre-defined highly efficient transpiration barrier fairly independent from rapidly changing environmental influences. Roots, in most conditions facing a soil environment with relative humidities very close to 100%, are orders of magnitude more permeable to water than leaf cuticles. Upon desiccation, the permanent wilting point of plants is defined as -1.5 MPa, which still corresponds to nearly 99% relative humidity in soil. Thus, the main reason for plant water stress leading to dehydration is the inability of root tissues to decrease their internal water potential to values more negative than -1.5 MPa and not the lack of a transport barrier for water in roots and leaves. Taken together, the commonly mentioned concepts that a drought-induced increase of cuticular wax or root suberin considerably strengthens the apoplastic leaf or root transport barriers and thus aids in water conservation appears highly questionable.