polymerase chain reaction (PCR), laboratory process in which a particular DNA segment from a mixture of DNA chains is rapidly replicated, producing a large, readily analyzed sample of a piece of DNA; the process is sometimes called DNA amplification.

The Process

In PCR, DNA (see nucleic acid) is immersed in a solution containing the enzyme DNA polymerase, unattached nucleotide bases (the subunits that DNA is composed of), and "primers," short sequences of nucleotides designed to bind with an end of the desired DNA segment. Two primers are used: one primer binds at one end of the desired segment on one of the two paired DNA strands, and the other primer binds at the other end but on the other strand. The solution is heated to break the bonds between the strands of the DNA. When the solution cools, the primers bind to the separated strands, and DNA polymerase quickly builds a new strand by joining the free nucleotide bases to the primers. When this process is repeated, a strand that was formed with one primer binds to the other primer, resulting in a new strand that is restricted solely to the desired segment. Thus the region of DNA between the primers is selectively replicated. Further repetitions of the process can produce billions of copies of a small piece of DNA in several hours.

Development and Applications

PCR was developed in 1985 by Kary B. Mullis, who was awarded the 1993 Nobel Prize in chemistry for his work. It is used in DNA fingerprinting and in medical tests to identify diseases from the infectious agent's DNA. In forensic use, the test can be used to compare two samples of DNA, usually by looking at matches (or mismatches) of six inherited traits (e.g., hair curliness) from each of the samples. Each trait is controlled by a single gene, each gene having at least two forms, or alleles, resulting in 21 combinations of these alleles, some of them very rare. A nonmatch conclusively excludes a suspect. PCR also is used in taxonomic classification to help show evolutionary relationships between organisms on the molecular level. It has the advantage of being able to be used even when only very small samples, such as tiny pieces of preserved tissue from extinct animals, are available.

open-chain compound: see aliphatic compound.

golden chain: see laburnum.

food chain: see ecology.

chain store: see store.

chain snake: see king snake.

chain reaction, self-sustaining reaction that, once started, continues without further outside influence. Proper conditions for a chain reaction depend not only on various external factors, such as temperature, but also on the quantity and shape of the substance undergoing the reaction. A chain reaction can be of various types, but nuclear chain reactions are the best known. A line of dominoes falling after the first one has been pushed is an example of a mechanical chain reaction; a pile of wood burning after it has been kindled is an example of a chemical chain reaction. In the latter case each piece of wood, as it burns, must release enough heat to raise nearby pieces to the kindling point. The wood, therefore, must be piled close enough together so that not too much heat is lost to the surrounding air. The conditions for a nuclear chain reaction can be understood by analogy. In the case of the fission of a nucleus, the reaction is begun by the absorption of a slow neutron. Each fission produces two or three fast neutrons. In order to sustain a chain reaction, a sample must be large enough to slow the neutrons so that one can be captured by another nucleus and produce a second fission. The sample must also be compact to prevent neutrons from escaping. The minimum quantity of a fissionable material necessary to sustain a nuclear chain reaction is called the critical mass. In a nuclear fission bomb, a chain reaction is started by forcing together two or more samples of fissionable material, each of less than critical mass, to form one sample of supercritical mass. The number of subsequent fissions produced by a single fission is always greater than one. The total number of fissions increases rapidly (exponentially) with time. In a fission reactor, the number of subsequent fissions for each fission must be exactly one. If the rate is less, the chain reaction will stop; if greater, it will soon grow out of control. In one type of fission reactor, a combination of fuel rods and control rods is moved in or out of a solid block of moderating material to control the reaction rate. In another type of reactor, the temperature of a liquid moderator controls the reaction. See also nuclear energy; nuclear reactor.

chain gang: see convict labor.

chain compound: see aliphatic compound.

chain, flexible series of connected links used in various ways, especially for the transmission of motive power, for hoisting (see pulley), and for securing or fastening. Commonly, mechanical energy from a motor or other source applied to a sprocket wheel is conveyed by means of an endless chain to another sprocket wheel for driving a mechanism. Examples of such an arrangement are found in bicycles, motorcycles, and conveyor belts. The chain in this application is so designed that each consecutive link fits over a sprocket, the distance between links being called the pitch. The relative speed of the wheels varies according to their relative circumferences and, thus, the number of sprockets on each. There are several types of chain for the transmission of power. A detachable-link chain has links that are simple rectangles, each with a connecting hook at one end by which it is attached to the next link. A pintle chain has links that are approximately U-shaped. The closed end of each link fits into the open end of the next one; a pin holds the two links together. A block chain consists of metal blocks that are joined together by side plates and pins to form links. A roller chain has links consisting of side plates with hollow cylindrical rollers between them. Pins pass through the rollers and side plates to hold the links together. A silent, or inverted-tooth, chain has links made of toothed metal plates. A number of these links are placed side by side to form a group. Each group is joined to another one by meshing the ends of the links of both groups and inserting a pin there. By repeating the process a chain can be formed. Its width can be varied by varying the number of links in a group. Although not completely silent, this type of chain is quieter than other power transmission chains. The coil chains used in hoists and for locking or fastening purposes are of the open-link type, comprising solid interlocked rings, or of the stud-link type, in which a stud, or bar, across the link keeps the chain from kinking.

Ratak Chain: see Marshall Islands.

Ralik Chain: see Marshall Islands.

Chain, Ernst Boris, 1906-79, English biochemist, b. Berlin, Germany. In 1933 he left Germany and went to England, where he conducted research at Cambridge from 1933 to 1935 and at Oxford from 1935; he lectured (1936-48) in chemical pathology at Oxford. In 1951 he became director of the International Research Center for Chemical Microbiology, Istituto Superiore de Sanità, Rome. He was professor of biochemistry at the Univ. of London from 1961. For his work on penicillin, Chain shared with Sir Alexander Fleming and Sir Howard Florey the 1945 Nobel Prize in Physiology or Medicine.

Laboratory technique used to make numerous copies of specific DNA segments quickly and accurately. These are needed for various experiments and procedures in molecular biology, forensic analysis (DNA fingerprinting), evolutionary biology (to amplify DNA fragments found in ancient specimens), and medicine (to diagnose genetic disease or detect low viral counts). Invented by Kary Mullis, PCR requires a DNA template (as little as one molecule) to copy, nucleotides to build the copies, and the enzyme DNA polymerase to catalyze the formation of bonds between the nucleotide monomers. Each three-step cycle (separating the two strands of the DNA double helix, marking the ends of the segment to be copied, and catalyzing the formation of bonds), which takes only minutes to complete, doubles the number of DNA strands present in the reaction medium. Repetition of this cycle many times results in an exponential increase in the amount of DNA.

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Sequence of transfer of matter and energy from organism to organism in the form of food. These interconnected feeding relationships intertwine locally into a food web because most organisms consume or are consumed by more than one other type of organism. Plants and other photosynthetic organisms (such as phytoplankton), which convert solar energy to food, are the primary food source. In a predator chain, a plant-eating animal is eaten by a larger animal. In a parasite chain (see parasitism), a smaller organism consumes part of a larger host and may itself be parasitized by even smaller organisms. In a saprophytic chain, microorganisms live on dead organic matter. Because energy, in the form of heat, is lost at each step, or trophic level, chains do not normally encompass more than four or five trophic levels.

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Process yielding products that initiate further processes of the same kind. Nuclear chain reactions are a series of nuclear fissions initiated by neutrons produced in a preceding fission. A critical mass, large enough to allow more than one fission-produced neutron to be captured, is necessary for the chain reaction to be self-sustaining. Uncontrolled chain reactions, as in an atomic bomb, occur when large numbers of neutrons are present and the reactions multiply very quickly. Nuclear reactors control their reactions through the careful distribution of the fissionable material and insertion of neutron-absorbing materials.

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or mail

Form of body armour worn by European knights and other medieval warriors. An early form, made by sewing iron rings to fabric or leather, was worn in late Roman times and may have originated in Asia. Medieval armourers interlaced the rings, which were closed by welding or riveting. In the 8th century, mail was a short coat with a separate sleeve for the sword arm. By the Norman Conquest (1066), the coat was long and fully sleeved; a hood, usually fitting under a helmet, covered the head and neck. By the 12th century, mail was fitted to hands, feet, and legs. The addition of plates to increase chest and back protection gradually evolved in the 14th century into complete plate armour, displacing mail.

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