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Immunologic Studies of Autochthonous Cancer an Evaluation of Several Procedures*

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Immunologic Studies of Autochthonous Cancer an Evaluation of Several Procedures* Immunologic Studies of Autochthonous Cancer An Evaluation of Several Procedures* ABRAHAM G. OSLER (Department of Microbiology, The Johns tlol~ldns Univers4ly School of Medicine and School q[ Hygiene and Public Health, and the Department of Medicine, The Johns Ilopkins University School of Medicine, Baltimore, Md.) INTRODUCTION sembling acquired immune tolerance or immuno- The unique and complex problems in studies of logic paralysis intervenes. spontaneous tumors of animals derive from the ~. Tumor-specific antibody is produced but is fact that the limiting frame of reference comprises not amenable to extensive in vitro investigation, a host response to an invasive, replicating cell pre- because it is of the cell-dependent type, which sumably of autologous origin. Unlike the situation mediates delayed hypersensitivity reactions, or of in many infectious diseases, immunologic phe- the reaginie, skin-sensitizing variety. nomena attributable to reactants of exogenous 3. Circulating tumor-specific antibody is syn- origin are not considered admissible in studies of thesized but is not uniformly detectable in the autochthonous cancer. As in the area of auto-anti- serum because it is bound to the cells which incite body production, this limiting condition imposes its production. rigorous requirements for the immunologist. Evi- 4. Antibody is synthesized by the tumor cells dence for an immune response to spontaneous tu- and is directed against host tissue components. mors must be buttressed by the demonstration of 5. Circulating tumor-specific antibody is pro- antigenic activity which clearly differentiates the dueed which reacts extensively with a variety of neoplasm from normal host tissues. normal tissue components. The recurrent reports and disavowals of spe- The reInainder of this report will be based on cific antibody production to autologous cancer tis- the assumption that circulating antibody, en- sues have led to the consideration of several hy- dowed with varying degrees of tumor specificity, is potheses. They are listed, not in the sense of being present in sera of humans or animals bearing spon- all-inclusive, and without further comment as to taneous tumors. Several experimental applications their validity, since a critical evaluation has not to the problem will be discussed primarily in terms yet been undertaken and the probability exists of the interpretational restrictions imposed by the that a single mechanism may not be equally ten- methodology. able for all situations. A. COMPLEMENT FIXATION IN ANTI- 1. Tumor cell multiplication progresses in the GEN CHARACTERIZATION STUDIES absence of antibody production. a) The genetic changes which characterize neo- 1. ThE FIvE-UNIT METHOD IN STUDIES plastic cells are not reflected in antigenic differ- OF TUMOR-SPECIFIC ANTIGENS ences, morphologic and functional disparities not- The complement (C') fixation reaction has been withstanding. used extensively for the immunologic characteriza- b) Tumor cells are deficient in antigenic com- tion of a tissue lipide with potential tumor spec- ponents present in normal tissues. ificity (1~, 44). It may therefore be appropriate to c) Tumor cells proliferate only when a state re- consider the applicability of current C' fixation * The studies carried out in the author's laboratory have procedures to this problem. The investigator who been supported by the National Science Foundation and the contemplates this approach faces the choice be- National Institute of Allergy and Infectious Diseases, United tween one of two general types of C ' fixation tech- Public Health Service. Financial assistance was also available nics: the conventional method which utilizes five from the Office of The Surgeon General, Department of the Army, under the auspices of the Commission on Cutaneous 50 per cent units of C' (C'Hs0) such as described in Diseases of the Armed Forces Epidemiological Board. (38) and the so-called quantitative technic with 50 1187 Downloaded from cancerres.aacrjournals.org on September 25, 2021. © 1961 American Association for Cancer Research. 1188 Cancer Research Vol. ~1, October 1961 or more CtH~0. Certain more general aspects of the the reaction may be localized between the limits of two procedures have been evaluated previously 4.5 to 5.5 units, since the range of partial lysis re- (~0, 33). For the present purposes, it may be ap- quires 0.5 to 1.5 C'Hs0. propriate to discuss other features with specific c) Complete lysis.--Less than 4.5 CtH~0 were applicational reference to the estimation and char- fixed. acterization of lipide-soluble tissue extracts from It is thus readily apparent that the use of 6 normal or cancerous tissues. C'Hs0 leads to a quantitative estimate only within The technic used by Rapport and Graf in their a narrow range of C' action--namely, one C'Hs0. studies of tumor lipides is essentially that devel- The remainder of the C' serves to swamp various oped by Wadsworth, Maltaner, and Maltaner (54) anticomplementary effects which may be very and also studied extensively by Rice (~7, 48). The troublesome in tests with less than 5 or 6 C'Hs0. A results are expressed in terms of the quantity of consequence of this severe restriction in the range antigen or antibody required to yield 50 per cent of C' action is that heavy reliance must necessarily hemolysis in a C' fixation reaction with 6 C~Hs0, be placed upon the differences in test results with and a constant amount of antiserum or antigen in the twofold dilution sequences of antigen or anti- the presence of varying quantities of the other. body. It follows, then, that an antigenic variable which is expressed by less than a 50 or 100 per cent difference could be attributable to "experimental Z error," which is of the same order of magnitude in / n... these reactions. i- I / z . I / / Ld / A simple illustration may clarify this point. The identification by Pangborn of cardiolipin as the o I / I / major antigenic constituent in crude lipide ex- ~a- I / I // tracts containing the Wassermann antigen (39, 40) '~\ /// w led to the isolation of a similar phospholipide from wheat germ, called sitolipin (5~). Comparative TYPE I serum titer estimations with these two phospho- I I I I lipides by conventional complement fixation and I 2 3 4 flocculation procedures supported the notion that RELATIVE ANTIGEN CONCENTRATION they were identical (46). 1 When quantitative CmaRT 1.--Two general types of C ~fixation reaction curves C' fixation technics with 50 C'Hs0 were applied to depicting the relative amounts of antigen and antibody re- this problem it was readily discernible that the quired to yield 50 per cent hemolysis (as adapted from [44]). phospholipide of plant origin was indeed similar but was not identical to those extracted from These, then, are two-dimensional block titrations, mammalian tissues. Sitolipin proved less efficient and the results are presented as curves drawn than the beef or human heart eardiolipins in pre- through the levels of immune reactants yielding 50 cipitating the Wassermann antibody or in C' fixa- per cent hemolysis as observed or interpolated tion studies with this antibody as derived from a values. The types of reaction curves that may be variety of sources including sera of human syphilit- obtained are given in Chart 1 as modified from ics (36, 37). (44). The data in Chart 1, which were obtained with The limitations of this method in studies of anti- the 6-unit method, describe the C' fixation results gen characterization may be summarized as fol- in the three different zones of antigen-antibody lows. interaction. The vertical line essentially parallel to When 6 units of C' are used in the initial fixation the ordinate roughly describes the region of anti- reaction, the subsequent addition of sensitized body" excess, becoming curved at equivalence zone erythroeytes to the antigen-antibody-C' mixture, ratios and asymptotic to the abscissa in the region for the lyric estimations, can lead to one of three of antigen excess. At antigen excess ratios, a di- results and implications, assuming that all the vergence may occur in that some immune systems controls are in good order. a) No lysis.--This finding means that at least exhibit more pronounced inhibition with excess 5.5 of the initial 6 C~Hs0 were consumed in reacting antigen (Type II) than do others (Type I). It is with the antigen, the antibody, or the specific ag- apparent from an inspection of these curves that gregate, since 0.5 C'Hs0 produces less than 10 per major segments of each type of curve contribute cent hemolysis. A. G. Osier, J. H. Strauss, and B. Lowenstein, unpublished b) Partial lysis.--The number of C'Hs0 fixed in observations. Downloaded from cancerres.aacrjournals.org on September 25, 2021. © 1961 American Association for Cancer Research. OSLER--Immunologic Studies of A utochthonous Cancer 1189 little to the characterization of the antigen-anti- diolipin in aqueous solution can combine with body reaction under study. With a Type I curve, Wassermann antibody but does not fix C' in this marked variations in antigen or antibody concen- interaction unless cholesterol is added. The choles- trations produce no detectable change in the C' terol is considered to facilitate aggregation by pro- fixation pattern. It is only in the restricted cur- viding a particulate surface for the adsorption of vilinear portion of this curve that alteration in the cardiolipin, whose unit molecular weight approxi- amount of either antigen or antibody will seriously mates e000 or less. The tendency for cholesterol affect the outcome of the reaction. These consider- crystals to aggregate spontaneously in isotonic ations may account for the wide variations encoun- saline must also be controlled with judicious addi- tered in those experiments which "did not permit tion of lecithin in carefully calibrated quantities. the quantity (of antigen) to be assessed accu- This cardiolipin-cholesterol-lecithin complex may rately" despite "corrections for the fixability of C' then be used in studies of antigenic activity based in terms of the deviation of the standard antigen on aggregation with antibody.
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