pollen grain surface is covered with waxes and proteins, which are held in place by structures called sculpture elements on the surface of the grain. The outer pollen wall, which prevents the pollen grain from shrinking and crushing the genetic material during desiccation, is composed of two layers. These two layers are the tectum and the foot layer, which is just above the intine. The tectum and foot layer are separated by a region called the columella, which is composed of strengthening rods. The outer wall is constructed with a resistant biopolymer called sporopollenin. Pollen apertures are regions of the pollen wall that may involve exine thinning or a significant reduction in exine thickness. They allow shrinking and swelling of the grain caused by changes in moisture content. The process of shrinking the grain is called harmomegathy. Elongated apertures or furrows in the pollen grain are called colpi (singular: colpus) or sulci (singular: sulcus). Apertures that are more circular are called pores. Colpi, sulci and pores are major features in the identification of classes of pollen. Pollen may be referred to as inaperturate (apertures absent) or aperturate (apertures present). The aperture may have a lid (operculum), hence is described as operculate. However the term inaperturate covers a wide range of morphological types, such as functionally inaperturate (cryptoaperturate) and omniaperturate. Inaperaturate pollen grains often have thin walls, which facilitates pollen tube germination at any position. Terms such as uniaperturate and triaperturate refer to the number of apertures present (one and three respectively). Spiraperturate refers to one or more apertures being spirally shaped. The orientation of furrows (relative to the original tetrad of microspores) classifies the pollen as sulcate or colpate. Sulcate pollen has a furrow across the middle of what was the outer face when the pollen grain was in its tetrad. If the pollen has only a single sulcus, it is described as monosulcate, has two sulci, as bisulcate, or more, as polysulcate. Colpate pollen has furrows other than across the middle of the outer faces, and similarly may be described as polycolpate if more than two. Syncolpate pollen grains have two or more colpi that are fused at the ends. Eudicots have pollen with three colpi (tricolpate) or with shapes that are evolutionarily derived from tricolpate pollen. The evolutionary trend in plants has been from monosulcate to polycolpate or polyporate pollen. Additionally, gymnosperm pollen grains often have air bladders,