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Translation Series No Original title: Morfologicheskie izmeneniia v tkaniakh ryb vokrug lichnok nekotorykh paraziticheskikh chervei.

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- 4 ARCHIVES THE LIBRARY FISHERIES REZCZCii 10.IRD OF CANADA NANAIA40, 5'gû FISHERIES RESEARCH BOARD OF CANADA Translation Series No. 580 Morphological changes in the fish tissue surrounding the larvae
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- 4 ARCHIVES THE LIBRARY FISHERIES REZCZCii 10.IRD OF CANADA NANAIA40, 5'gû FISHERIES RESEARCH BOARD OF CANADA Translation Series No. 580 Morphological changes in the fish tissue surrounding the larvae of certain parasitic worms By I. G. Mikhailova, E. V. Prazdinkov, and T. O. Prusevich Original title: Morfologicheskie izmeneniia v tkaniakh ryb vokrug lichnok nekotorykh paraziticheskikh chervei. From: Trudy Murmanskogo Morskogo Biologicheskogo Instituta, Vol. 5, No. 9, pp Translated by G. N. Kulikovsky, Bureau for Translations, Foreign Language Division, Department of the Secretary of State of Canada Fisheries Research Board of Canada Biological Station, Nanaimo, B. C. 1965 Morphological Changes in Fish Tissue around 251 the Larvae of some Parasitic Worms. By I.G. Mikhailova, E.V. Prazdnikov, and T.O. Prusevich. (Laboratory of Comparative and Experimental Embryology. Chief - B.P. Tokin.) (From Trudy Murmanskogo Morskogo Biologicheskogo Institute /iransaction of the Murmansk Sea-Biological institute/, USSR Academy of Sciences, No. 5 (9) p ). The problem formulated in the Thirties by E.N. Pavlovsky: MDrganism as the medium of habitation (1934, 1937, 1945) acquires an ever-increasing actuality. As a point on the agenda the necessity remains to mobilize the assistance of other Sciences for its solution:- of Experimental Zoology, General Pathology, Biochemistry, Immunology, and of Microbiology. New ways of considering the interrelations between the parasite and the host also appear on the borderline between parasitology and embryology. A successful method of study of the problem of the immunity of the embryos (Tokin, 1955a, 1955b) is primarily a study of the problem of embryology - a problem of interrelation of at least two organisms: of a macroorganism and microorganism. In the light of ideas developed by B.P. Tokin, the interrelation between the parasite and the host, in particular between the host and the larval stages of the parasites, deserve special attention, because: a parasite, which in the course of evolution has found itself in another body, which is to the former, both a medium of habitation and food, cannot be merely an enemy to the host. With a certain foundation, one may speak conditionally about the host as being the enemy, and furthermore, - a very dangerous enemy to the parasite (Tokin, 1955a, page 10). Being in possession of the knowledge concerning the 2/ immunological properties of the host and of the parasite, one could, by increasing the defensive reaction of the host and by decreasing the immunological reaction of the parasite, annihilate the later. The facts, concerning the destruction, incapsulation, or phagocytosis of parasites inside the body of the host, are commonplace. Nevertheless, we still know next to nothing about the causes of the death of a parasite.- in some cases, and of a normal development and preservation in the tissues of the host, - in other cases. It is not accidental that, when discussing the problems of interrelations between the host and the parasite, the parasitologists do not fail to note the necessity for taking the immunological features into account (Shuurmans-Stekhoven, 1955; Schultz, Davtyan, 1954, 1955; Shikhobalova, 1950; Naumova, 1955; Bauer, 1958; Markov, 1958, and others). At the same time, the morphological studies of the defensive reactions of the tissues of the host and of the parasite are fen, and they are either descriptions of individual casuistic cases (Heim, 1954; Tratnig-Frankl, 1956), or a supplement to the parasitological or zoological observations of the authors. Extremely controversial ideas were developed by V.A. Fausek (1903, 1913), who tried to draw a direct analogy between parasitism and viviparity. However, Fausek must be credited with being the first to give special attention to the utilization, by the parasitic organism, of the inflammation processes in the tissues of the host for the purpose of feeding. In the work of R.M. Lewert and Lee Chang-Ling (1954) interesting 252 data concerning histochemical changes in the host's tissues around the larvae of the parasitic worms are presented. There are detailed descriptions in the literature of various interrelations occurring between the tissues of the host and the larval stages of the parasitic worms, primarily in 1.1ch intermediate hosts as mollusks (Agersborg, 1924; Rees F.G., 1934; Rees W.J. 1936, and others). In a work by E.A. Zelikman Certain ecological-parasitical connections on the littoral of the northern portion of the Kandalaksha Bay (1955), one may find rather complete accounts of bibliographical data available on this problem. E.A. Zelikman, just as other scientists, observed the destruction picture of 3/252. the visceral complex of the mollusks in the development of the sporocysts and rediae. The author describes cases of antagonistic action of one parasite upon another at double invasion. In the works of E.D. Logachev (1956, 1961), of Logachev and N.A. Fedyushkina (1955), of Logachev and B.R. Bruskin (1959), peculiar multinuclear structures are described, which originated in the tissues of a mollusk produced by a parasite. Efforts are made to establish, how the reparative processes. run in the parasitic worms. A large number of observations indicate that the character of the membranes and capsules, originating near the endoparasite, change considerably in case of a change in the life functions of the host or the parasite. When the parasite dies, there is either a resorption of the latter by means of the phagocytes of the host, or the remnants of the parasite are enclosed in a dense fibrous capsule, which is later permeated with calcium salts. The data concerning the interrelations of tissues of haddock and cod, with the larvae of nematoda selected by us for the experiment (Anisakis sp., Contracoecum aduncum, Terranova decipiens) and of the cystoda (Pyramico- cephalus phocarum, Bothriocephalus sp.) may be found in the works of a number of researchers. A.Ya. Bazikalova (1932) points out the pathological importance of parasites and the reactive changes owing to their influence in the organs of the host. Thus the plerocercoids (Cestodes gen. sp.) produce considerable changes in the liver of haddock and cod. The colour of the liver turns dark red, irnd the plerocercoids degenerating in the parenchyma are surrounded by voluminous hardenings of irregular shape. Bazikalova writes that similar formations, something like cases of smaller size, are also formed around the larvae of the parasitic nematodes Anisakis sp., Contracoecum aduncum. total weight of the parasites in the Gadidae, according to the observations of the author, does not exceed hundredths or thousandths of one per cent of the total weight of the body of the fish. The larvae of the Cestodes produce a more destructive effect upon the tissues of the host, than do the larvae of the nematodes, particularly when localized in the liver. Bazikalova believes that the basic pathogenous effect is supplied by the toxines of the parasites, these toxines produce partial destruction The 4/ and regeneration of the host's tissues. The weight of the haddocks and cod's liver decreases with the increase of the number of parasites, the fattiness also decreases (the relation between the weight of the liver and the weight of the fish). The author came to the conclusion, that parallel with other aspects of the study of the problem of the character of the effect of the parasites upon the host's body, the histological changes, particularly those originating as the result of the affection of the liver by the parasites, deserve special attention. S.S. Shulman (1948, 1958), R.E. Shulman (1950), S.S. Shulman and R.E. Shulman-Albova (1953) record the infestation of the liver of the Gadidae by Contracoecum aduncum and by Anisakis sp. The weight of the liver decreases in the affected fish. The most distinct decrease in the weight of the liver presents itself in the largest specimens (haddock and cod between 1 and kilograms). The celozoic parasites in the Novaya Zemlya char (Dogel and Markov, 1937) change in connection with the migration from fresh to sea water. The autumn chars are not infested with Bothriocephalus, and the summer chars are not infested with Contracoecum. This phenomena, together with some very interesting observations by a number of authors, including Polyansky (1955), concerning the decrease, in the winter time, of the infestation rate of cod and haddock with celozoic and intestinal parasitic nematodes (Contracoecum and Anisakis), still remain unexplained. Thus, despite the agreement of the majority of the parasitologists on the importance of the morphological studies of the reactive changes originating in the organs of animals infested with endoparasites, this side of the problem remains poorly studied. The aim of the present paper is the study of the morphological characteristics of the capsules around the larvae of certain parasitic worms in the tissues of fish, mainly of haddock (Melanogrammus aeglifinus), cod (Gadus morhua morhua), shorthorn sculpin (Myoxocephalus scorpius), and also partly of the arctic char (Salvelinus alpinus). Capsules were studied in the liver and the mesentery of the haddock, cod, sculpin, and char. The length of the studied fish was: haddock and cod centimeters, sculpin centimeters, and char centimeters. 5/253 Organ parts with larvae of parasitic worms were prepared in Bouin solution, in zenker-formol, Carnoy's solution, and in 10% formalin. The best results were obtained from the treatment with Bouin and Carnoy.'p.: solutions. Fragments were poured over by celloidin-parafin and by celloidin. The sections were stained by the Mallory method and with resorbin-fuchsin by the van Gison method; they were silvered according to Foot's method; stained by means of Hansen's trioxihematein, and also with picro-fuchsin. In the liver of haddock and cod, and also of the shorthorn sculpin one may find a considerable number of larvae of Anisakis sp., Contracoecum aduncum, and less frequently of Terranova decipiens. Detailed analysis of the intensity of the infection of this organ in the haddock, cod, and sculpin may be also found in the works of A.Ya. Bazikalova (1932), S.S. Shulman (1948), S.S. Shulman and R.E. Shulman-Albova (1953), and Yu.I. Polyansky (1955). In the present paper we describe the morphology of certain agechanges ir the capsules in late stages of development on the basis of histological analysis of 38 cases of the liver infection of the cod, haddock, and sculpin by the larvae àf the Nematodes, and in 8 cases - by the larvae of the Cestodes.. Bodies of the anisakid larvae in the vast majority of cases are encapsulated and wound up in spiral form. In the parenchyma of the liver, in places where the parasite is located, one observes impressions, the 'depth of which corresponds, as a rule, to the thickness of the capsule, which repeats the windings of the larval body. It should be noted, that when the parasite stays longer, these impressions are expressed more sharply. The formation () I f the capsule, obviously, takes place in the following manner. The larvae of Anisakis sp. and Contracoecum aduncum attach themselves to the surface of th'e liver, thus they compress and mechanically damage the serous membrane and the adjoining portions of the liver parenchyma. Besides the mechanical factor, the products of the life of the larvae obviously also have effect upon the adjoining tissues. All this leads to the reactive hypertrophic growth of the fibroblastic elements and to the accumulation of the cells of hematogeneous origin. The insignificant distrophic changes in the adjoining liver cells also take place simultaneously. The juxtaposition of the thickness of the relatively normal serous membrane of the liver of the haddock and the and the relatively still young capsule of the connective tissue, 6/ appears extremely demonstrative (fig. 1, 2). The serous membrane near the larva becomes tens of times thicker. The age of the capsule in each observation case remains unknown 254 to us, but the studied capsule may be morphologically distinguished into two groups: the young and the old . This division is also supported by the experimental data on the study of the formation rates of the capsules (Prusevich, in the present compendium). Morphological changes occurring in the serous membrane and in the adjoining sections of the liver parenchyma, when the larvae of Anisakis sp. and of Contracoecum aduncum are present, are similar in many respects, - in both cases a capsule of connective tissue develops near the larvae, at the expense of the hypertrophic changes in the serous membrane of the liver. Larvae of the parasitic nematodes plunge into the glandular tissue of the liver (fig. 3) and produce insignificant local degenerative changes. We shall not dwell on the differences in the reactivity of the liver tissues of the haddock, cod and sculpin, when responding to the invasion of parasitic larvae, because we observed comparatively monotype changes in the connective tissues of this organ and in the liver parenchyma, except in a few cases, when we observed in the liver of cod extensive infiltration of tissues of the capsule by the leucocytes. The morphology of the capsules around the larvae of the Anisakis sp., and of the Contracoecum aduncum does not differ in principle. We present here a description of the capsules, which originate around the larvae of Anisakis sp., since we possess considerable amount of material on the development of capsules of this species. By histological analysis one is able to distinguish in a young capsule formed near the larva of Anisakis sp., conditionally and morphologically, at least three layers of connective tissue. The first layer which directly contacts Anisakis sp., is a narrow strip of tissue consisting of damaged cellular elements with traces of pyknosis, and sometimes of the lysis of the nuclei. Here the collagen fibrills are homogenized and in spots the entire basic material of the connective tissue is intensively tinted with acid stains (fig. 3, 4, and 5). 256 Behind the layer of damaged tissue follows a second layer with a greater quantity of fibroblastic elements, these are partly subjected to 11.1 7/ degenerative changes. The nuclei of the cells of this layer are light coloured, sometimes bacilliform, often unusually elongated (fig. 5). It is known, that the nuclei of the fibroblasts of the mammals take such a shape when damaged. Pictures of fragmentation and amitotic division of the nuclei of the fibroblasts are often observed. Collagen fibril's occur among the cellular elements of this layer. The third layer usually consists of porous connective tissue (fig. 4). It contains numerous blood vessels, diverse cellular elements, among which often occur histiocytes, large fibroblasts with a clearly pronounced diplasmatic differentiation. Amitotically dividing fibrocytes are visible in certain preparations. Among the cell elements there frequently occur leucocytes, extra-vascular erythrocytes, and remnants of destroyed erythrocytes, which are like pigment grains. In places where the thin connective-tissue lacerti which encase the body of the larva depart, original widenings occur, at the base of which one often observes blood or lymphatic vessels. At the border of the liver parenchyma there is a relatively unchanged more solid connective tissue, which reminds us of an externally normal serous membrane. Individual elastin fibres are observed in the third layer, but they are very few in number, and probably do not have any significance in the formation of the capsules. Possibly, this is connected with the fact that the larvae of Anisakis sp. are immobile in the fish tissues. The liver tissue adjoining the connective-tissue capsule of the larva is very little changed campared to the other part of this organ, except for a stronger development of the connective-tissue interlayers and except for their more numerous content of blood vessels. The basic connective-tissue skeleton of the cod and goby liver 257 are the argyrophyl tissues and granules, which are particularly numerous along the course of the capillaries. Thin collagenous fibrills occur along the course of the blood vessels, and are sometimes observed between the cells of the liver. The most numerous collagenous bundles are located near the large blood vessels and in the serous membrane of the capsule. The argyrophil fibres were studied inside the capsules near the larvae of the nematodes in the liver of cod and sculpin. In both cases their analogous arrangement was observed. Individual argyrophil granules occur in the first layer. In the second layer we see numerous small argyrophil granules, sometimes the argyrophil substance is deposited near the nuclei of the degenerating cells. 8/ The argyrophil granules are most numerous of the capsules of the nematode larvae in the sculpin, where the second layer is sharply distinguished from the third layer, because of the presence of fine argyrophil granularity in the second layer. In the third layer argyrophil fibres occur, these form retiform structures in some areas. At the border with the second layer, in a number of cases there originates an interlacing of argyrophil and collagen fibrills. At the border with the parenchyma of the liver, the argyrophil fibres arrange themselves along the periphery of blood vessels. With the increase of the period of stay of the Anisakis sp. in the serous membrane of the liver, the connective tissue grows in the adjoining sections. An increasing number of blood vessels penetrate into the capsule. As a result of intra-abdominal pressure, and, probably, also because of the local deceleration in the growth of the liver tissue, the nematode together with connective-tissue capsule plunges deeper into the liver parenchyma. Along the course of the blood vessels the thickness of the connective tissue interlayers increases, and a part of the blood vessels penetrate from the parenchyma into the capsule. The connective-tissue casing , which repeats the helices of the larva, appears distinctly in the histological sections. In older capsules the morphology of individual layers is somewhat different. The basic substance of the connective tissue appears in the first interior layer, it is homogeneously tinted over with orange-yellow colour. The cell elements disappear from this layer. Between the first and the second layer (the layer of the fibroblastic elements) appears a narrow strip of tissue, which does not contain cell elements, which, according to Mallory, is tinted in lilac hues, and which sometimes shows fibrillar structures. In the second layer one observes the destruction of a part of the cells. This layer has no distinct borders and blends with the first layer, and is probably 258 permeated with calcium salts. The third layer is the widest one; two zones are observed in this layer (fig. 6). In the first zone, which adjoins the second layer, we find blood vessels. In the connective tissue degenerative cell elements, the more or less numerous errant cells, are observed (fig. 6, 7). In preparation stained with picro-fuchsin, the homogenized collagen bundles are seen in this zone. poorly stained with acid fuchsin. substance is obviously formed. Sometimes the basic substance is homogeneous and is In these cases, the homogenized collagen In the second zone, at the border to the 9/ liver parenchyma, wide collagen bundles are arranged. In cod and haddock these bundles are like intertwining ribbons, while in the shorthorn sculpin collagen window-like membranes appear. At the described stage of development, the capsules separate easily from the serous membrane of the liver. To clarify the formation rate of the capsules around the larvae of parasitic nematodes, we have carried out tests on transplanting of larvae of Anisakis sp. from cod to sculpin. To
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