Trichuris Ovis (Abildgaard, 1795) (Fam
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STUDIES ON THE LIFE-HISTORY OF TRICHURIS OVIS (ABILDGAARD, 1795) (FAM. TRICHURIDAE: NEMA, TODA) BY GOBIND SINGH THAI'AR,M.Sc., PH.D. (LOND.), F.N.I., F.A.Sc., F.Z.S.I. AND KR. SURESH SINGH, PH.D. (LUCK.) (From the Department of Zoology, Universityof Lucknow) Received on March 8, 1954 Trichuris ovis (Abildgaard, 1795) is a very common parasite of the domesti- cated animals and is found not only in the c~ecum of sheep and goats in India but has also been reported from cattle and buffaloes. A light infection usually does not cause much damage but sometimes acute infections cause colitis, chronic diarrhoea and other disorders of the intestinal tract. It is also possible that these worms may cause trauma and irritation of the intestinal tract. A large number of workers have studied the life-history of the different species of Trichuris. The earliest account is by Davaine (1858) who for the first time showed that the eggs of T. trichiurus pass out of the host unseg- mented and require about 34 weeks to embryonate. Leuckart (1876) working on T. ovis demonstrated that, the life-history is simple and takes place by direct infection with embryonated eggs. He successfully repeated the experi- ments on swine by using eggs of T. crenatus. Railliet (1884), while working with T. vulpis, showed that only the embryonated eggs are infective, and non-embryonated eggs pass out without development. Grassi (1887) [quoted by Faust (1939)] for the first time described the stages in the life- history of T. trichiurus. Neshi (1918) [quoted by Yokogawa (1920)], working on T. vulpis in dogs, said that the larva~ migrate through the lungs as they do in ascarids. Tochihara (1922) working on T. vulpis in dogs, and F[illeborn (1923) working on T. leporis in rabbits, found that no migration through the lungs takes place. Hasegawa (1924), working on T. vul'pis in dogs, found that the larva~ in the early stages migrate to the crypts of Lieberkuhn. Brown (1927) studied the rate of development of eggs in T. trichiurus under field conditions and found that eggs do not embryonate in direct sunlight. Spindler (1929) found that eggs of T. vulpis exposed to air at 77~o saturation at 22 ~ C. died in 12 days and m 69 70 GOBIND SINGH THAPAR AND KR. SURESH SINGH that there is a direct correlation between rainfall and geographical distribu- tion of human trichurids. Hkasawa (1929) working on T. vulpis and T. trichiurus did not find any evidence of the larvae reaching or migrating through the lungs. Nolf (1932) found that freezing temperature is lethal to the eggs of human trichurids. Matsubayashi (1932), working on 7". vulpis in rats, followed the local migration of the larvae in the host's intestine. Artjueh (1936) concluded that I". ovis enters through the crypts of Lieberkuhn and that they are blood feeders. Dinnik (1937) found that eggs of T. tri- chiurus start dividing 25-30 hours after passing out of the host and also gave the nuclear changes occurring during the early development of the eggs. Dinnik (1937, 1937 a, 1939) studied the development of the eggs of T. tri- chiurus under different environmental conditions and found that the egg shell can resist watery solutions of copper sulphate, hydrochloric and nitric acids, mercuric chloride and formalin. He also observed that the shell resists desiccation by an exchange of water through the polar plugs. These eggs can do without oxygen for several days during their development. Miller (1939) studied the development and hatching of eggs of T. vulpis. Bonilla Naar (1945) gave an account of the life-history of T. trichiurus. Shikhobalova and Gorodilova (1945) found that eggs of T. trichiurus require an average temperature of 33-35 ~ C. and a minimum soil humidity of 8~o for development. Miller (1947) worked out in detail, the early phases of the life-history of T. vulpis in dogs, and further confirmed the observations of Hasegawa 0924) in finding the migration of the early stages in the crypts of Lieberkuhn. Deo (1948, 1950) infected kids with embryonated eggs of T. ovis and recovered adult worms after 136 and 161 days respectively, but he gave no structural details. It would thus appear that no attempt has been made to give a complete and connected account of the life-history of this parasite. In a closely allied genus, Capillaria, divergent views have been expressed about its life-history. Ftilleborn (1924) showed that the larvae of C. hepatica (syn. Hepaticola hepatica) migrate to the liver through the hepatic portal system and this was confirmed by Nishigori (1925), who also gave a detailed account of its life-history. When the infected liver is accidentally eaten by another host or the food is contaminated (Momma, 1930; Troisier and Deschiens, 1930; Shorb, 1931), the larvm hatch in the intestine and migrate to the liver through the hepatic portal system and sometimes reach lung, brain, kidney or skin. Vogel (1930) showed that young larvae of C. hepatica, if implanted on other organs, reach back liver. Luttermoser (1938) found that the eggs of C. hepatica withstand desiccation and freezing for 1-2 weeks and about 10~o of eggs remain viable after 28 weeks of encapsulation in Life-History of Trichuris ovis (Abildgaard, 1795) 71 liver, and also the larwe penetrate the c~cum wall 6 hours after infection and reach liver via the hepatic portal system (Luttermoser, 1938 a). In some other species of Capillaria, the life-history may be simple and direct, as in C. columbce, C. contorta and C. cerophila (Cram, 1932, 1936; Wehr, 1937; Levine, 1938; Christenson, 1938) or it may be requiring an inter- mediate host, as in C. caudinflata, C. annulata, C. plica and C. mucronata, for which species of earthworms are true intermediate hosts (Wehr, 1936; Allen and Wehr, 1942; Morehouse, 1944; Petrov and Borovkova, 1942; Zucchero, 1942; Wehr and Allen, 1945; Skarbilovich, 1945; Allen, 1949). In view of these divergent views about the life-histories of the members of the family Trichurid~e and also regarding differences of opinions about migration of the larvae through the body of the host even in Trichuris species, it was considered desirable to reinvestigate the life-history and the methods of infection in T. ovis. The results indicate that there is neither migration through the body of the host nor is there an intermediate host involved in the life-cycle of this parasite. In the present work it is attempted to study the life-history of the parasite under laboratory conditions and experiments conducted to infect goats and other small laboratory animals with a view to ascertain the time taken to complete the life-history under varying conditions. Since the details of the early stages with special reference to migration have been studied by Miller (1947), greater attention had been paid in this communication to the later stages of the life-history. The present work was carried out under a grant from the Indian Council of Agricultural Research, New Delhi. Our thanks are due to the authorities of the Council for providing funds for this work. We are also thankful to the authorities of the Lucknow University for providing laboratory facilities. MATERIALS AND METHODS Gravid females of T. ovis were obtained from the local slaughter-houses and the eggs for cultures were recovered from them by cutting the body into small pieces. For final infection laboratory-reared kids were used. These animals were kept under sanitary conditions and care was taken to avoid natural infection taking place in the animals. The earthworms for the infection experiments were reared from cocoons in the laboratory and were kept in sterile earth. They were exposed to infection according to the methods of Wehr (1936), Allen and Wehr (1942), More,house (1944) and Wehr and Allen (1945). BI* 72 GOBIND SINGH THAPAR AND KR. SURESH SINGH The eggs for cultures were obtained from females by teasing the uterus in a petri dish full of sterile distilled water. Care was taken to wash, as much of the debris from cultures as possible, to avoid infection by fungus and other organisms. The water of the culture was changed every day and hence there was no necessity to use dilute formalin to check the growth of fungi and bacteria. It was, however, observed that a slight infection did not hamper the development of the eggs in any way. The eggs were kept in a constant temperature cabinet at 80-82 ~ F. [NFECITION EXPERIMENTS To ascertain if an intermediate host is required in the life-cycle of T. ovis, earthworms, Pheretima posthuma, Eutyph~eus waltoni and Allolobophora sp. were exposed to embryonated eggs. These earthworms were examined at different intervals of time and all of them were found to be negative for larvae and hence it was concluded that there is no intermediate host required in the life-history of T. ovis. Two kids born under laboratory conditions were then selected for the final infection. These animals were not allowed to eat grass, etc., but were fed on goat's milk. The f~eces of both the animals were examined regularly and it was negative for helminthic infection till the experiment started. A definite number of embryonated eggs were then fed to the kids by medicinal dropper for a number of days. The following table gives the dates and the number of eggs fed to kid A. Experiment L Kid. A 3- 9-1946 .. approx. 200 eggs 4- 9-1946 .. ,, 200 ,, 5- 9-1946 .. ,, 200 ,, 2-10--1946.. ,, 300 ,, 3-10--1946.. ,, 300 ,, 4-10-1946 .