Antiidiotype antibodies carrying the internal image

The idiotypes of a lymphocytic receptor can be located on various segments of the variable regions and may require the contribution of both the heavy and light chain. Some idiotopes are associated with the combining site, since their interaction with antiidiotype antibodies can be inhibited by antigen. Other idiotypes are located on variable region segments which are not involved in the combining site; the interaction of these idiotypes, collectively designated the 'framework associated idiotypes', with anti-Id antibodies is not antigen inhibitable. Therefore, it appears that the clones recognizing idiotypes (Ab2) represent a heterogeneous population displaying various specificities.

These clones express surface immunoglobulin and secrete antibody which can be classified into four major categories (Figure 1):

1. Ab2a, which is specific for idiotypes associated with the framework segments of the variable region. The binding of Ab2a to Abl does not alter the binding of Abl to antigen. These antibodies can exhibit a regulatory effect, priming

Figure 1 Various types of anti-idiotype antibodies.

or suppressing Ab1 clones, but they cannot replace an antigenetic stimulus. It was shown that allogeneic A/J anti-T15Id antibodies, which are not antigen inhibitable, can induce a long-lasting suppression of T15 anti-phosphocholine antibodies. Similarly, a monoclonal antibod) specific for a cross-reactive idiotype shared by antibodies specific for influenza virus neuraminidase was able to prime animals for a specific rcsponse but not elicit by itself anti-neuramini-dase antibodies.

2. Ab2y are antibodies that recognize combining site-associated idiotypes. Such antibodies can inhibit the binding of Abl to antigen but cannot elicit an Abl response when injected into an animal. Such antibodies can have a regulatory effect, despite the fact that they are devoid of internal antigen properties.

3. Ab2(3 are anti-idiotypic antibodies mimicking the antigen. Such anti-Id antibodies bind to the combining site of the lymphocytic receptor and are able to stimulate the expansion of clones and their maturation in antibody-forming cells. The interaction of Ab2|3 with the combining site of Ab l is obligatorily antigen inhibitable, since Ab2 idiotype is an antigen surrogate.

4. Ab2e (epibody) are antibodies which bind both to idiotype and to an epitope of self or foreign antigen.

The major questions concerning the stimulatory effect of anti-Id antibodies are related to:

• The ability to distinguish Ab2a or Ab2y from Ab2p.

• The determination of the immunogenic and tolerogenic doses. If the idiotypes are surrogates of the antigens, then they may obey the same rules of immunogenicity as the antigens.

• The requirements of MHC-restricted recognition of internal image antigen.

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