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Secondary antibody antigen reactions occur as a result of antigen multivalency, and result in agglutination or precipitation of a polymeric antigen antibody network. A visible product is formed over a time scale of minutes to hours (Eq. 5.2):
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5:2
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Both primary and secondary Ag Ab interactions may be exploited for analytical purposes, but quantitative bioanalytical methods make extensive use of primary binding interactions. The intrinsic association constant that characterizes antibody epitope(hapten) binding is called the af nity: ! Ab H Ab : H Affinity K Ab : H = Ab H 5:3 5:4
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Typical af nity values7 for antibody hapten interactions range from 105 to H 1012 M 1 . Because of the very high af nities involved, the reversal of Ab binding may be accomplished only under extreme conditions: at high or low pH values, at high ionic strength, or in the presence of chaotropic ions (e.g., perchlorate) or denaturants (urea, guanidinium) that interfere with hydrogen bonding. Af nity values are experimentally determined by maintaining a constant total Ab concentration, and varying the H concentration. Quantitation of H free and H bound then allows the construction of a Scatchard plot, where H bound = f H free Ab total g is plotted against H bound = Ab total . The slope of this plot is the negative of the af nity, K. Scatchard plots for monoclonal and polyclonal antibodies are shown in Figure 5.4. Note that polyclonal antibodies yield a curved Scatchard plot due to their range of epitope selectivities and af nities.8
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Figure 5.4. Scatchard plots for monoclonal and polyclonal antibodies.
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ANALYTICAL APPLICATIONS OF SECONDARY ANTIBODY ANTIGEN INTERACTIONS
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For polyclonal antibodies, experimental K values represent an average af nity, which is usually represented as K0 . 5.5. ANALYTICAL APPLICATIONS OF SECONDARY ANTIBODY ANTIGEN INTERACTIONS 5.5.1. Agglutination Reactions Agglutination occurs when an antibody interacts with a multivalent, particulate antigen (i.e., an insoluble particle), resulting in cross-linking of the antigen particles by the antibody. This eventually leads to clumping, or agglutination, and will not occur with haptens. Agglutination may occur when a unideterminate, multivalent antigen (i.e., possessing many copies of single epitope A ) interacts with a single antibody (Anti-A); it may also occur if a multideterminate, univalent antigen (with epitopes A, B, C, etc.) interacts with at least two distinct antibodies (Anti-A plus Anti-B, e.g.). Whether or not agglutination occurs depends on the relative concentrations of antigens and antibodies. The agglutination test is used to qualitatively test for the presence of antibodies in serum, and is also used in blood grouping. As an example, we will consider a test for the presence of antibodies to the bacterium Brucella abortus.9 In this test, a series of test tubes are prepared, each containing the same quantity of the bacterial suspension, which is the antigen. To each tube is added increasingly dilute solutions of serum. If the antibody is present, agglutination will be observed over a range of serum dilutions, as shown in Figure 5.5. Two dilution zones are apparent, one called the prozone and the other, the agglutination zone . In the prozone, no difference is apparent after serum is added and allowed to incubate for several minutes. In the agglutination zone, a visible precipitate forms, and settles to the bottom of the tube. The high-dilution end of the agglutination zone is called the titer of the serum, and is used as a semiquantitative expression of the Ab concentration for the comparison of sera. Note that tests made on sera at only one dilution may give misleading results, because of the lack of agglutination in the prozone and at high serum dilutions.
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Figure 5.5. Agglutination test for Brucella abortus antibodies.9 [Reprinted, with permission, from E. Benjamini, G. Sunshine, and S. Leskowitz, Immunology: A Short Course, 3rd ed., Wiley-Liss, New York, 1996, pp. 115 118. ISBN 0-471-59791-0. Copyright # 1996 by Willey-Liss, Inc.]
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