Red Blood Cells
Red blood cells are the most common type of blood cell and the vertebrate body’s principal means of delivering oxygen from the lungs to body tissues via the blood.
Red blood cells are also known as RBCs or erythrocytes (from Greek erythros for “red” and kytos for “hollow”, with cyte nowadays translated as “cell”). A schistocyte is a red blood cell undergoing fragmentation, or a fragmented part of a red blood cell.
Erythrocytes consist mainly of hemoglobin, a complex molecule containing heme groups whose iron atoms temporarily link to oxygen molecules in the lungs or gills and release them throughout the body. Oxygen can easily diffuse through the red blood cell’s cell membrane. Hemoglobin also carries some of the waste product carbon dioxide back from the tissues. (In humans, less than 2% of the total oxygen, and most of the carbon dioxide, is held in solution in the blood plasma). A related compound, myoglobin, acts to store oxygen in muscle cells.
The color of erythrocytes is due to the heme group of hemoglobin. The blood plasma alone is straw-colored, but the red blood cells change color depending on the state of the hemoglobin: when combined with oxygen the resulting oxyhemoglobin is scarlet, and when oxygen has been released the resulting deoxyhemoglobin is darker, appearing bluish through the blood vessel walls. Pulse oximetry takes advantage of this color change to directly measure the arterial blood oxygen saturation using colorimetric techniques.
The sequestration of oxygen-carrying proteins inside cells (rather than having them dissolved in body fluid) was an important step in the evolution of vertebrates; it allows for less viscous blood and higher concentrations of oxygen.
Erythrocytes in mammals are anucleate when mature, meaning that they lack a cell nucleus and thus have no DNA. In comparison, the erythrocytes of nearly all other vertebrates have nuclei; the only known exception is salamanders of the Batrachoseps genus. Mammalian erythrocytes also lose their other organelles including their mitochondria and produce energy by fermentation, via glycolysis of glucose followed by lactic acid production. Furthermore, red cells do not have an insulin receptor and thus glucose uptake is not regulated by insulin. As a result of the lack of nucleus and organelles, the cells cannot produce new structural or repair proteins or enzymes and their lifespan is limited.
Mammalian erythrocytes are biconcave disks: flattened and depressed in the center, with a dumbbell-shaped cross section. This shape (as well as the loss of organelles and nucleus) optimizes the cell for the exchange of oxygen with its surroundings. The cells are flexible so as to fit through tiny capillaries, where they release their oxygen load. Erythrocytes are circular, except in the camel family Camelidae, where they are oval.
In large blood vessels, red blood cells sometimes occur as a stack, flat side next to flat side. This is known as rouleaux formation, and it occurs more often if the levels of certain serum proteins are elevated, as for instance during inflammation.
The spleen acts as a reservoir of red blood cells, but this effect is somewhat limited in humans. In some other mammals such as dogs and horses, the spleen sequesters large numbers of red blood cells which are dumped into the blood during times of exertion stress, yielding a higher oxygen transport capacity.
The diameter of a typical human erythrocyte disk is 6–8 µm, much smaller than most other human cells. A typical erythrocyte contains about 270 million hemoglobin molecules, with each carrying four heme groups.
Adult humans have roughly 2–3 × 1013 red blood cells at any given time (women have about 4 to 5 million erythrocytes per microliter (cubic millimeter) of blood and men about 5 to 6 million; people living at high altitudes with low oxygen tension will have more). Red blood cells are thus much more common than the other blood particles: There are about 4,000–11,000 white blood cells and about 150,000–400,000 platelets in each microliter of human blood. The red blood cells store collectively about 3.5 grams of iron, more than five times the iron stored by all the other tissues combined.
The process by which red blood cells are produced is called erythropoiesis. Erythrocytes are continuously being produced in the red bone marrow of large bones, at a rate of about 2 million per second. (In the embryo, the liver is the main site of red blood cell production.) The production can be stimulated by the hormone erythropoietin (EPO), synthesised by the kidney; which is used for doping in sports. Just before and after leaving the bone marrow, they are known as reticulocytes which comprise about 1% of circulating red blood cells. Erythrocytes develop from stem cells through reticuloctyes to mature erythrocytes in about 7 days and live a total of about 120 days. The aging cells swell up to a sphere-like shape and are engulfed by phagocytes, destroyed and their materials are released into the blood. The main sites of destruction are the liver and the spleen. The heme constituent of hemoglobin is eventually excreted as bilirubin.