HEMOGLOBIN

also spelled Haemoglobin, protein in the blood of many animals-in the red blood cells (erythrocytes) of vertebrates-that transports oxygen to the tissues. Hemoglobin forms an unstable, reversible bond with oxygen; in the oxygenated state it is called oxyhemoglobin and is bright red; in the reduced state it is purple-blue. Asphyxiation occurs when the blood contains an excess of substances (e.g., carbon monoxide) with which hemoglobin forms semipermanent or permanent compounds instead of reversible bonds with oxygen. Hemoglobin develops in cells in the bone marrow that become red blood cells. When red cells die, hemoglobin is broken up: iron is salvaged, transported to the bone marrow by proteins called transferrins, and used again in the production of new red blood cells; the remainder of the hemoglobin forms the basis of a chemical called bilirubin that is excreted into the bile, eventually to reach the intestine, where its end product gives the feces their characteristic yellow-brown colour. Each hemoglobin molecule is made up of four heme groups surrounding a globin group, forming a tetrahedral structure. Heme, which accounts for only 4 percent of the weight of the molecule, contains all the iron and gives a red colour to the molecule. Globin consists of two linked pairs of polypeptide chains. The development of each chain is controlled at a separate genetic locus. The amino acid sequences in these chains have been fully worked out; single or multiple substitutions along the chains result in abnormal hemoglobins. The study of abnormal hemoglobins has provided a considerable amount of information on human evolution and history. Hemoglobin S, for example, is found in cases of sickle-cell anemia, a severe, hereditary form of anemia in which the cells become crescent-shaped when oxygen is lacking (see sickle-cell anemia). The sickling trait is found almost exclusively in blacks of the Negroid geographic race, among whom, in certain parts of Africa, it may reach high frequencies. Other hemoglobin variants occur in various parts of the world and help scholars to trace past human migrations and to study genetic relationships among contiguous and separated populations. See also erythrocyte.