 "The waxy sheet of cuticle also functions in defense, forming a physical barrier that resists penetration by virus particles, bacterial cells, and the spores and growing filaments of fungi".The plant cuticle is one of a series of innovations, together with stomata, xylem and phloem and intercellular spaces in stem and later leaf mesophyll tissue, that plants evolved more than 450 million years ago during the transition between life in water and life on land.Together, these features enabled upright plant shoots exploring aerial environments to conserve water by internalising the gas exchange surfaces, enclosing them in a waterproof membrane and providing a variable-aperture control mechanism, the stomatal guard cells, which regulate the rates of transpiration and CO2 exchange.