Aleurone (from Greek aleuron, flour) is a protein found in protein granules of maturing seeds and tubers. The term also describes one of the two major cell types of the endosperm, the aleurone layer. The aleurone layer is the outermost layer of the endosperm, followed by the inner starchy endosperm. This layer of cells is sometimes referred to as the peripheral endosperm. It lies between the pericarp and the hyaline layer of the endosperm. Unlike the cells of the starchy endosperm, aleurone cells remain alive at maturity. The ploidy of the aleurone is (3n) [as a result of double fertilization]. Aleurone protein Aleurone proteins can have two different morphological features, homogenous and heterogeneous. The homogenous aleurone consists of similar protein bodies (e.g. Phaseolus vulgaris) while the heterogeneous aleurone consists of granules of different shapes and types of proteins covered with a membrane (e.g. Ricinus communis). Aleurone tissue The aleurone layer surrounds the endosperm tissue of grass seeds and is morphologically and biochemically distinct from it. Starchy endosperm cells are large, irregularly shaped cells and contain starch grains while aleurone cells are cuboidal in shape and contain aleurone grains. In most cultivated cereals (wheat species, rye, oats, rice and maize) the aleurone is single-layered, whereas barley has a multicellular aleurone layer. Thick primary cell walls enclose and protect the aleurone cells. The aleurone layer is important for both the developing seed and the mature plant. The aleurone tissue accumulates large quantities of oils and lipids that are useful during seed development. It is also a site of mineral storage and in some species, functions in seed dormancy. The aleurone may also express several pathogen-protective proteins including PR-4. Aleurone also serves as the most dietarily beneficial fraction in many brans. In addition, the aleurone tissue contains many protein-storing vacuoles known as protein bodies. In cereals with starchy endosperm, the aleurone contains about 30% of the kernel's proteins. In multicolored corn, anthocyanin pigments in the aleurone layer give the kernels a dark, bluish-black color. Aleurone development The development of the aleurone layer involves several periclinal, and anticlinal cell divisions and several steps of genetic regulation. The dek1 gene and crinkly4 (cr4) kinase both function as positive regulators of aleurone cell fate. The normal dek1 gene is needed in order to receive and respond to positional cues that determine the fate of aleurone cells during development. Mutants of the dek1 gene block the formation