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Tumors in the Invertebrates: a Review BERTASCHARRER*ANDMARGARETSZABÓLOCHHEAD

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Tumors in the Invertebrates: a Review BERTASCHARRER*ANDMARGARETSZABÓLOCHHEAD Tumors in the Invertebrates: A Review BERTASCHARRER*ANDMARGARETSZABÓLOCHHEAD (From the Department of Anatomy, University of Colorado School of Medicine, the Department of ZniHogy, ?'nirer.tity of Vermont, and the Marine Biological Laboratory, Woods Hole, Massachusetts) INTRODUCTION papers in which invertebrate tumors are reviewed Tumors are the result of abnormal cell prolifera to some extent are two in French (13, 148) and tion. Therefore, the study of tissue growth, normal one in Russian (30). and abnormal, constitutes the central problem of A. critical survey of the present status of the tumor research which thus becomes essentially a problem of invertebrate tumors encounters serious biological problem. When approached from this difficulties. For one thing, the «latainthe literature broader point of view, an analysis of tumorous are often controversial, and many of the descrip growth should include representatives from all tions are inadequate or hard to evaluate. Patholo- groups of living organisms. It has been recognized gists specializing in tumor research are not, as a that the study of plant tumors yields significant rule, familiar with invertebrate material. On the results. Within the animal kingdom comparative other hand, zoologists, versed in the intricacies of pathology has concerned itself largely with neo invertebrate anatomy and taxonomy, are usually plasms in various groups of vertebrates (74, 10!), inexperienced in the diagnosis of tumor growth. 112, 124), while invertebrates have been all but Furthermore, the terminology developed almost neglected. As a matter of fact, until fairly recently exclusively for use in mammalian pathology invertebrate tissues were often considered in should not be applied to invertebrate animals, capable of developing tumorous growths. until the analogies between vertebrate and in Teutschlaender (144) believed that tumors can vertebrate tissues are more thoroughly under not occur in animals at a phylogenetic level lower stood. than the fishes. Engel (25) elaborated extensively In spite of such difficulties two facts stand out on the reasons why invertebrates are unable to as the result of this survey: invertebrate tissues are develop cancer. Emphasizing anatomical differ capable of tumorous change, and they offer an op portunity to approach the study of tumors in new- ences between invertebrates and vertebrates he discussed chiefly three points. The first two con ways. cern differences in the nervous and vascular sys TUMORS IN VARIOUS GROUPS tems; the third deals with the embryonal theory of OF INVERTEBRATES tumor growth. Believing that invertebrate cells, on The first question one may ask is :In which types account of their considerable regenerative power, of invertebrates have tumors or cellular reactions are embryonic in character, Engel concluded that comparable to tumorous growth been observed? they cannot revert to the embryonic stage and The information available refers to almost all the produce tumors. In the light of modern biological major invertebrate phyla. Tumors are said to oc concepts these views are obsolete, not only with cur in annelids, sipunculids, arthropods, molluscs, regard to anatomical considerations, but also he- and ascidians. Even among the most primitive cause they are based on inadequate material. metazoans—dicyemids (91)—and among the pro Actually, during the past .50 years a consider tozoans (82) nuclear anomalies have been observed able literature on spontaneous and experimentally which are comparable to atypical mitoses as found induced tumors in invertebrates has grown. In in certain mammalian neoplasms. The best known addition, numerous observations exist concerning group and the most interesting from the point of various tissue reactions that may be more or less view of comparative oncology are the insects. closely related to tumorous growth. These data are scattered and often not easily accessible. The only VARIOUS TISSUE REACTIONS AND * Special Research Fellow, I'.S. Public Health Service. Re TUMORLIKE GROWTHS search grant»fromthe American Cancer Society and The Anna Great caution is indicated in an evaluation of Kuller Ktiiul are gratefully acknowledged. the types of tumors occurring in invertebrate ani Received for publication, April 14, 1950. mals. For one thing, the difficulty of arriving at a 40.-5 Downloaded from cancerres.aacrjournals.org on September 25, 2021. © 1950 American Association for Cancer Research. 404 Cancer Research satisfactory classification, which exists even in tumorous changes is the effect of certain parasites mammalian pathology, is multiplied in inverte on the so-called fat body of insects, a type of con brates. Furthermore, in a number of instances a nective tissue which stores reserve materials and, variety of formations in invertebrates which can in certain species at least, shows no sign of cellu not be considered as true tumors have been desig lar proliferation after metamorphosis. Under the nated as such. Among these are cysts, often the influence of the parasites these adipose cells have result of what is called an "inflammatory process." been observed to hypertrophy and to resume mi- It would be preferable to call such processes "in totic activity in the adult stage. Many of these jury reaction" or "repair hyperplasia," since ac mitotic figures are abnormal (67, 79, 106). The cording to Menkin (78) the term inflammation, result is a hyptrplasia of the fat body of these in which implies vascular response, should be re sects (16, 17, 18, 23, 99). stricted to vertebrates. A frequently observed situ Another instance where caution in the u.se of ation is the following (e.g., 8, 11, 24, 81, 54, 56, 59, terms is indicated is that of the insect "myce- 60, 61, 64, 68, 69, 80, 83, 98): almost any disturb tomes" (structures caused by intracellular sym- ing factor (foreign body, parasite, tissue implant biotes). The practice of Mahdihassan (75, 76) of with or without bacterial contamination, degener considering these biologically useful structures of ating organ, hereditary tissue anomaly, sterile bacterial origin as tumors is not in keeping with agar or celloidin, or irritant, such as croton oil) will commonly accepted views (135, chapters 4 and (i: cause an accumulation of lymphocytes in the af 137, chapter 5). Likewise, parasitic structures such fected area. In many invertebrates certain blood as those described by Mclntosh (77) in the caudal cells may change from free-moving to sessile ele region of Sagitta (Chaetognatha) are not tumors ments and vice versa under various conditions (27, in the strict sense of the word. 28, 73, 163). When they accumulate, often in im Many of these pathological formations are evi pressive numbers in the vicinity of irritants, dently not true tumors but may show certain re phagocytosis and encapsulation of the center of semblances to neoplastic growth. Thus, it is ob disturbance take place. In the cysts thus formed vious that in many instances the decision as to the cells making up the capsule may undergo whether or not a reported structure is to be classi changes, sometimes of a degenerative nature. Such fied as a neoplasm meets with considerable dif- tissue responses resulting in well defined "growths" culty. A great deal of further observation is neces may easily be mistaken for tumorous processes. In sary before an opinion can be formulated. Just as most cases they merely indicate an injury reac certain tumors in vertebrates are considered to be tion, but it is not unlikely that under certain con on the borderline between hyperplasia and benign ditions such cysts may eventually give rise to tumors, some of the formations reported here in tumor-like growths or real tumors. the invertebrates might prove to be borderline Various abnormal growths, with or without ul cases. cération, have been observed on the body wall of Nevertheless, so far as can be judged from the molluscs collected in nature or reared in the labora available evidence, cases of true tumors, both be tory. The causes of these growths are unknown. nign and malignant, appear to occur among in In each case the histológica! picture showed the vertebrates. These cases are discussed in the fol presence of an injury reaction with repair hyper lowing section, in which the term "tumor" will be plasia, characterized by densely packed nuclei of freely used wherever the balance of evidence seems connective tissue cells, fibroblasta, and numerous to suggest a true neoplastic growth. Also discussed migrating cells. Tissue reactions experimentally in this section will be certain experimental work of produced by burning and by strong acid were es relevant interest, including cases in which no neo sentially the same as these natural growths (139, plastic growth was obtained. 141, 142). Another type of cellular response in inverte CAUSES OF INVERTEBRATE TUMORS AND TUMOR-LIKE STRUCTURES brates which in itself should not be considered as tumorous growth manifests itself in the hypertro Spontaneous occurrence.—A number of cases re phy of cells due to the action of parasites. For ex ported in the literature concern incidental obser ample, in the intestinal epithelium of certain vations of so-called spontaneous tumors. It is per marine worms gregarines may cause up to a ten haps more accurate to classify them as tumors fold increase in cellular size accompanied by whose cause is unknown. Among the earliest ob changes in the appearance of nuclei and cytoplasm servations is one on a lobster (Homarus) by Mc lntosh who "many years ago, described a tumour (9, 10, 102, 136, 145). Somewhat more significant in connection with which originated in the wall of the grinding storn- Downloaded from cancerres.aacrjournals.org on September 25, 2021. © 1950 American Association for Cancer Research. SCHARRERAND LOCHHEAD—Tumors in Invertebrate* 405 ach and pushed its way through the carapace be cur which are derived from several smaller hind the eyes. The tumour enlarged and finally re cells by cytoplasmic, and subsequently nuclear, sulted in the death of the lobster, which was a very fusion (20).
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