Proteins are large organic compounds made of amino acids arranged in a linear chain and joined together by peptide bonds between the carboxyl and amino groups of adjacent amino acid residues. The sequence of amino acids in a protein is defined by a gene and encoded in the genetic code. Although this genetic code specifies 20 "standard" amino acids, the residues in a protein are often chemically altered in post-translational modification: either before the protein can function in the cell, or as part of control mechanisms. Proteins can also work together to achieve a particular function, and they often associate to form stable complexes.
Like other biological macromolecules such as polysaccharides and nucleic acids, proteins are essential parts of organisms and participate in every process within cells. Many proteins are enzymes that catalyze biochemical reactions, and are vital to metabolism. Proteins also have structural or mechanical functions, such as actin and myosin in muscle, and the proteins in the cytoskeleton, which forms a system of scaffolding that maintains cell shape. Other proteins are important in cell signaling, immune responses, cell adhesion, and the cell cycle. Protein is also necessary in animals' diets, since they cannot synthesise all the amino acids and must obtain essential amino acids from food. Through the process of digestion, animals break down ingested protein into free amino acids that are then used in metabolism.
The word protein comes from the Greek πρώτα ("prota"), meaning "of primary importance" and these molecules were first described and named by the Swedish chemist Jöns Jakob Berzelius in 1838. However, proteins' central role in living organisms was not fully appreciated until 1926, when James B. Sumner showed that the enzyme urease was a protein. The first protein to be sequenced was insulin, by Frederick Sanger, who won the Nobel Prize for this achievement in 1958. The first protein structures to be solved included hemoglobin and myoglobin, by Max Perutz and Sir John Cowdery Kendrew, respectively, in 1958. Both proteins' three-dimensional structures were first determined by x-ray diffraction analysis; the structures of myoglobin and hemoglobin won the 1962 Nobel Prize in Chemistry for their discoverers.