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Significant Biochemical Effects of Hepatocarcinogens in the Rat: a Review* E

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Significant Biochemical Effects of Hepatocarcinogens in the Rat: a Review* E Significant Biochemical Effects of Hepatocarcinogens in the Rat: A Review* E. REID (Chester Beally Research Instthde, institute of Cancer Research: Royal Cancer Hospital, London, S.W. 3, England) CONTENTS INTRODUCTION Introduction In precancerous liver, as obtained by giving Validity of comparing “hepatomas―with nor rats a hepatocarcinogen for several weeks, there ma! liver are diverse biochemical abnormalities. These are Validity of comparing precancerous liver with now surveyed and appraised with respect to their normal liver or with hepatomas possible significance for neoplasia. Consideration Bases for expressing results is also given to primary hepatomas, but only Present Approach briefly to transplanted hepatomas. In the vast Rate-limiting steps literature on hepatocarcinogenesis, there have been Trial of various hepatocarcinogens and of in diverse speculations, but only a few attempts active analogs (e.g., 53, 83, 149, @10)to collate and weigh the Trial of treatments that potentiate or retard evidence. hepatocarcinogenesis In the study of neoplastic changes by biochemi Survey of Biochemical Data cal as distinct from histochemical methods, the Does precancerous liver show biochemical ab use of liver has obvious advantages—notably the normalities similar to those in primary hepa predominance of one type of cell. Parenchymal tomas? cells comprise ca. 90 per cent of the weight of “Score-card―for significance of observed ab the liver in the rat, although only about 65 per normalities cent of the actual number of cells (46, 80, 196). Might any of the abnormalities entail rate Nevertheless, there are certain assumptions and changes in metabolic processes? pitfalls that have not always been recognized. The reversibility criterion Validity of comparing “hepatomas―withnormal Search for hepatomas with minima! deviations liver.—Uncritical comparisons have often been from normal liver. Dramatic deviations made between normal liver tissue and “hepa Conclusions tomas,― repeatedly transplanted and conceivably Theories of hepatocarcinogenesis unlike the original tumor (83, 149, 151), or primary Future work liver tumors which may have been bile-duct tu References mors, or even hyperplastic nodules. Transplanted hepatomas, or hepatomas cultured in intro or * The experiments in the reviewer's laboratory' were sup @ by grants to the Chester Beatty Research Institute (In rendered ascitic, can of course give valuable in stitute of Cancer Research: Royal Cancer Hospital) from the formation (149); but neglect to study primary Medical Research Council, the British Empire Cancer Cam hepatomas is hardly excusable merely because paign, the Anna Fuller Fund, and the National Cancer Insti of the time and work needed to induce the tumor tute of the National Institutes of Health, U.S. Public Health Service. Biochemical differences among transplanted The following abbreviations are used: AAF, @-acetylamino hepatomas, or between transplants and primary fluorene (@-fiuorenylacetamide); AAT, o-aminoazotoluene; hepatomas, are briefly considered later. In at AB, aminoazobenzene (not azobenzene); MAE, 4-methyl-AB; least one study (@)where transplanted hepatomas DAB, 4-dimethyl-AB (N,N-dimethyl-p-phenylazoaniline); DMN, dimethylnitrosamine; DNA, deoxyribonucleic acid; differed biochemically from primary tumors, the RNA, ribonucleic acid. Conventional abbreviations are used for latter may have been of bile-duct origin (“choli nucleotides—e.g., AMP, adenosine-5'-monophosphate; GDP, angiomas,― sometimes termed “choliangiocarci guanosine-5'-diphosphate; UTP, uridine-5'-triphosphate; d (as nomas―). In comparison with primary hepatomas, prefix), deoxy. primary choliangiomas may be lower in alkaline Received for publication September @1,1961. phosphatase (69), in rhodanese (17@), in xanthine 398 Downloaded from cancerres.aacrjournals.org on September 24, 2021. © 1962 American Association for Cancer Research. REID—BiOChemiCal Effects of Hepatocarcinogens 399 oxidase ([1@], but see [164]), and in the capacity fair agreement on the following categories (69, to synthesize serum albumin (3@); but higher in 153, 170, 195) : (a) trabecular hepatoma, small the capacity to synthesize phospholipide (80). celled or large-celled (cf. 146) ; (b) adenocarcinoma; No marked differences have been found in gly (c)anaplasticcarcinoma;(d)mixedhepatomaand cçdysisrate (@6), in drug-metabolizing reactions choliangioma. Trabecular hepatomas are less ab (1), or in the content of ribonucleoprotein particles normal than other types with respect to $-hy (144). Hyperplastic nodules (“hyperplastomas,― droxybutyrate oxidation (61), although not with “regenerative foci―), perhaps premalignant (115, respect to the content in the supernatant fraction 119), resemble normal liver at least with respect of RNA and of acid ribonuclease.' In the latter to the oxidative capacity of the mitochondria respects small-celled hepatomas differ little from (60, 61). large-celled hepatomas,' but their glucose-6-phos Primary hepatomas themselves vary in histolo phatase and tryptophan peroxidase content may gy (69, 84). With 3'-Me-DAB, although not with be lower, and they may be morphologically more DAB, the proportion of “parenchymal―cellstends cognate to biliary epithelium (146). to be lower than that in normal liver (50, 53). It is then, important, that tumors as used for Lymphocyte infiltration, or changes such as re biochemical comparison with normal liver should generation, cirrhosis, fibrosis, necrosis, or cyst indeed be malignant and be hepatomas in the formation (53), may complicate biochemical stud sense of having histological affinities to liver pa ies and be irreve!ant to the process of carcinogene renchyma. The latter stipulation implies a com sis (69, 105, 119). Cirrhosis may contribute to the promise between two extreme points of view. It low catalase activity of hepatomas (1@) and has been suggested that even supposedly liver-cell necrosis to the high free ATPase (166) and low tumors originate from bile-duct cells, proliferation rhodanese (7@) activities—although not to the of which may indeed occur in precancerous liver low xanthine oxidase activity (164) or the ap (153). On the other hand it has been suggested parently high glycolysis (@6).However, acetanilide that even choliangiomas are of parenchymal origin acy!ase is low in hepatomas but somewhat in (105, 195) or, less improbably, that choliangiomas creased with cirrhosis (106), and nucleases are and liver-cell tumors have a common origin in a low in intact hepatoma cells but high in necrotic primitive hepatic cell (119). areas (53). Isotopic studies indicate that the tissue Male rats fed a 20 per cent protein diet con lying deep within a hepatoma nodule differs from taming 3'-Me-DAB (88) eventually develop highly that in an outer layer 5 mm. deep, probably be malignant tumors, which in our experience are cause of impaired circulation associated with ne mainly trabecular hepatomas or adenocarcinomas, crosis (215). It is, then, advisable to reject the and which are in the form of discrete nodules central part of the nodule. It is also useful to such that there is no excuse for including adjacent pass the tissue through a disk with perforations nonmalignant tissue in the tumor sample (cf. of under 1 mm. in diameter, thereby reducing the 68). 4'-F-DAB, in common with AAF (which content of fibrous and other nonhepatoma ele is rather slow-acting, especially in old rats [115]), ments (53, 156). is somewhat toxic. Dimethylaminobenzene-1-azo The study of metastases, from sites such as the 1-naphthalene (69) was only slightly carcinogenic lungs and diaphragm, may be advantageous on in our hands. DAB has been widely used but often histological grounds (170); but the rat may die with neglect of the possibility that the tumors before the metastases are large enough for bio were choliangiomas. With supplementary B vita chemical examination. In a comparison of primary mins (80), or with a low-protein diet (53,2)—which, hepatomas with metastases, the depression in cat however, could give protein-deprivation effects alase and N-demethylase activities was much less in the controls (110)—DAB can indeed produce marked with the metastases (122). hepatomas. Hepatomas used for biochemical studies should, Firminger (69) draws attention to the finding then, be routinely checked histologically for non of Opie that, in the absence of cirrhosis, the right hepatoma elements as a minimum precaution, lobe of the liver is particularly prone to develop and the presence of metastases should be noted tumors, presumably because most of the portal as one criterion of malignancy. To facilitate com blood goes to this lobe. Such a localization of parison of results from different laboratories, a tumors, although not found by the Millers, has description should be given of the gross and micro been seen in other laboratories; but biochemical scopic appearance, perhaps with a dassification such as medical pathologists employ. There is 1E. Reid, unpublished experiments. Downloaded from cancerres.aacrjournals.org on September 24, 2021. © 1962 American Association for Cancer Research. 400 Cancer Research Vol. @2,May 1962 changes have not been found earlier in the right showed an increase in bile-duct tissue to 5 per lobe than in the other 23 cent at 2 weeks and to 30 per cent at 4 weeks. It seems self-evident that controls should be Similar changes, expressed in terms of cell number, used with rats given no carcinogen but otherwise
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