Amebiasis: Infection and Disease by Entamoeba histolytica Editor Roberto R. Kretschmer Head, Division of Immunology Unidad de Investigacion Biomedica Instituto Mexicano del Seguro Social and Subdivision de Medicina Experimental Facultad de Medicina Universidad Nacional Autonoma de Mexico Mexico, D. F. CRC Press Taylor & Francis Group Boca Raton London New York CRC Press is an imprint of the Taylor & Francis Group, an informa business CRC Press Taylor & Francis Group 6000 Broken Sound Parkway NW, Suite 300 Boca Raton, FL 33487-2742 Reissued 2019 by CRC Press © 1990 by Taylor & Francis Group, LLC CRC Press is an imprint of Taylor & Francis Group, an Informa business No claim to original U.S. Government works This book contains information obtained from authentic and highly regarded sources. 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Disclaimer The publisher has made every effort to trace copyright holders and welcomes correspondence from those they have been unable to contact. ISBN 13: 978-0-367-24452-1 (hbk) ISBN 13: 978-0-429-28253-9 (ebk) Visit the Taylor & Francis Web site at http://www.taylorandfrancis.com and the CRC Press Web site at http://www.crcpress.com To Liliana, Verónica, and Kilian PREFACE There were mixed feelings when the invitation to write a book on amebiasis reached me in 1985. First, several excellent reviews, both in specialized journals and in books, had appeared recently or were in press.13 A. Martínez-Palomo, a leader in the field of amebiasis, had just contributed to re-establish the scientific relevance of this parasite — recently de- serving only chapter treatment in the Anglo-Saxon literature — by publishing a book totally devoted to the subject.4 He subsequently edited another book on amebiasis, where several authors rendered their ''state-of-the-art" account of a field that had witnessed a remarkable progress in the last two decades.5 Furthermore, there lurked in the background the imminent appearance of J. I. Ravdin's extensive text with 50 chapters covering virtually every aspect of this remarkable parasite.6 To make matters worse for me, most of the collaborators I could think of for such a new book on amebiasis had, not unexpectedly, contributed to one or the other of the aforementioned texts. After some internal and external deliberation, however, I decided to accept the challenge of presenting yet another medium-size book on amebiasis, provided two conditions could be met. In the first place, the enterprise would have be be postponed for at least 2 years in order to allow enough new information to accumulate. I thought this not to be an unreasonable proposition in a field of such remarkable vitality. Second — and foremost — the authors should be allowed, alas invited, not to shun controversy, and in fact engage in reasonable, constructive speculation, a salutary exercise in a field where so much and sometimes so conflicting information is at hand. Such an effort could help pave the way for future de- velopment by perhaps suggesting pathways for research. Having agreed upon these two premises, the chosen collaborators — all experts in their own field with immediate experience with amebas and their victims — responded positively to the invitation, and the signal to start was given in the summer of 1987. Most recent texts on amebiasis stress the importance of this disease as a worldwide health problem that has been surprisingly and systematically neglected by the international health agencies.7 Ranking diseases by importance can be a treacherous and thankless task.8 Still, is is obvious that amebiasis has not received its due priority i f one considers its prevalence (500,000,00 0 infection s pe r annum), its morbidity (50,000,00 0 diseases per annum ranging from trivial to life-threatening) an d its mortality (50,000 to 100,000 per annum).9 Thus, amebiasis still kills annually more people than AIDS has killed in the decade since its recognition (an estimated 75,000), yet receives only a very small fraction (ca. 10~3) of the financial suppor t deservedly allotte d to HIV research.10 Granted, amebiasi s is in principle a reasonably preventable and curable disease that has apparently reached a certain epidemiological stability affecting mostly underdeveloped countries. AIDS in contrast, though theoretically perhaps as preventable as amebiasis, is still an incurable disease that is undergo- ing an explosive epidemiological expansion, most alarmingly in rich, industrialized countries. Curiously, both diseases are converging in some individuals at risk with striking but as yet unpredictable long-term consequences.11 For physicians in developed countries, amebiasis may soon cease to be a forgotten problem affecting forgotten people.12 Moreover, AIDS is revealing the fallacy of strictly nationally oriented disease-control strategies. Or confront, if you wish, amebiasis with other leading parasitic diseases of mankind: malaria,8 with a very well-known mode of transmission (and thus potentially effective control) produces far more recurring illness and death than amebiasis, killing alone in Africa yearly ten times more people (mostly children) than amebiasis worldwide.13 Yet, among protozoarian diseases the death toll of amebiasis is only second to that of malaria.9 South American trypanosomiasis is way down in prevalence (12,000,000 infections per annum), but morbidity and mortality (60,000 deaths per annum) are only slightly below those of amebiasis. Giardiasis is perhaps as prevalent as amebiasis, but its morbidity — occasionally in epidemic bursts — is lower, and its mortality practically nil. As far as transmission is concerned, both diseases have much in common (fecalism), an d are therefore equally difficult t o control in daily life. Among the larger parasites (helminths), schistosomiasis is about half as prevalent and morbid (ca. 200,000,000 and 20,000,000 cases per annum respectively) as amebiasis but its mortality (1,000,000 deaths per annum) is much higher. Finally, ascaris, the giant intestinal round- worm, is the most prevalent infection of man (ca. 1,000,000,000 cases per annum) but has a relatively insignificant morbidity and only rarely causes death. As in the case of amebiasis, its control is difficult a s it would ultimately require massive, long-term improvements in sanitary and agricultural practices. Clearly, the priorities for disease control advocated by the international health agencies take into consideration not only prevalence, the age group mostly affected, morbidity, and mortality but, quite importantly, the feasibility of adequate and effective control as well.7 That explains, on the one hand, the high priority bestowed on malaria and the upgrading of schistosomiasis, and on the other hand the low priority assigned to ascariasis, giardiasis, and amebiasis, notwithstanding their high prevalences, and in the case of amebiasis also its relatively high mortality. In spite of its relatively high morbidity and mortality (only slightly lower than amebiasis), South American trypanosom- iasis (Chagas' disease) is also ranked low because of its difficult control. All this provides insight into the complex if not bizarre sociological interactions of ranking, promoting, lobbying, and funding that a given disease experiences on its way to public recognition. Through the efforts of the Rockefeller Institute, amebiasis has at least been identified as a pristine example in their Great Neglected Diseases Program.7 This timely strategic decision led to a multicentric renaissance in amebic research, reaping a generous harvest of challenging new knowledge. The late Bernardo Sepulveda was the undisputed pioneer of this awakening in Mexico, a country where E. histolytica is particularly at home. It is in his spirit that we embarked in this enterprise.14 The text is organized in 11 chapters. It follows the traditional distribution of first basic and then clinical material, preceded by a chapter on the history of amebiasis that adds some fascinating material on the Mexican contribution in this field of human knowledge. The chapter on Immunology wa s split into four subchapter s (antigens , humoral immune re- sponses, cellular immune responses, and effector events). Likewise, the chapter devoted to the clinical aspects of amebiasis was divided into two parts: one covering adults, the other children. To all collaborators I wish to express my gratitude for their enthusiastic response. Special thanks should go to our colleagues in Israel and in the U.S., as they had to suffer th e additional burden of slow communication with the editor, in a world where only bad news appears to travel fast and cheap. The encouragement and seasoned advice received from Dr. Ruy Pérez-Tamayo and Dr. Adolfo Martínez-Palomo is greatly appreciated. We all benefited from the translating and editorial skills of our Editorial Secretary, Ms. Isabel Pérez-Montfort, to whom our gratitude is hereby expressed. Finally, the writing of this book was supported by a generous grant (PCSACNA-451740- A) from the Mexican Council for Science and Technology (CONACYT). In doing so it recognized both the paramount importance of this disease on a national scale and the con- tributions made by Mexican scientist s to the understanding o f this cosmopolitan healt h problem. Roberto R. Kretschmer Ciudad Universitaria México, D.F. 1989 CONTRIBUTORS Ingeborg Becker Armando Isibasi Subdivisión de Medicina Experimental Head, Division of Immunochemistry Facultad de Medicina Unidad de Investigación Biomédica UNAM Instituto Mexicano del Seguro Social México, D.F., México México, D.F., México Bibiana Chavez Roberto R. Kretschmer Department of Experimental Pathology Head, Division of Immunology Centro de Investigación y Estudios Unidad de Investigación Biomédica Avanzados del IPN Instituto Mexicano del Seguro Social México, D.F., México and Subdivisión de Medicina Experimental Ann Chayen Facultad de Medicina Department of Biophysics UNAM Weizmann Institute México, D.F., México Rehovot, Israel Jesus Kumate Louis S. Diamond Secretary of Health Head, Section of Parasite Growth & Secretaría de Salud Differentiation México, D.F., México Laboratory of Parasitic Diseases NIAID National Institutes of Health Martha López-Osuna Bethesda, Maryland Division of Immunology Unidad de Investigación Biomédica Araceli Gonzalez Instituto Mexicano del Seguro Social Departamento de Inmunología México, D.F., México Instituto de Investigaciones Biomédicas UNAM México, D.F., México Adolfo Martinez-Palomo Head, Department of Experimental Arturo González-Robles Pathology Department of Experimental Pathology Centro de Investigación y Estudios Centro de Investigación y Estudios Avanzados del IPN Avanzados del IPN México, D.F., México México, D.F., México Marco Antonio Meraz Vicente Guarner Department of Cellular Biology Professor Centro de Investigación y Estudios Universidad Lasalle Avanzados del IPN Hospital Angeles del Pedregal México, D.F., México México, D.F., México Gonzalo Gutierrez Isaura Meza Director de Planeación, Head, Department of Cellular Biology Información y Evaluación Centro de Investigación y Estudios Secretaría de Salud Avanzados del IPN México, D.F., México México, D.F., México David Mirelman Ruy Pérez-Tamayo Professor Head, Subdivisión de Medicina Department of Biophysics Experimental Weizmann Institute Facultad de Medicina Rehovot, Israel UNAM México, D.F., México Irmgard Montfort Blanca Ruiz Subdivisión de Medicina Experimental Research Associate Facultad de Medicina Departamento de Biologia Molecular UNAM Instituto de Investigaciones Biomédicas México, D.F., México Facultad d e Medicina UNAM Onofre Muñoz México, D.F., México Director Hospital de Pediatría del Centro Médico Norberto Treviño García-Manzo Nacional, Instituto Mexicano dei Director General de los Servicios Seguro Social Médicos del Departamento del Distrito México, D.F., México Federal, México México, D.F., México Ruy Pérez-Montfort Senior Researcher Haydee K. Torres-Guerrero Departamento de Microbiología Department of Cellular Biology Instituto de Fisiología Celular Centro de Investigación y Estudios UNAM Avanzados del IPN México, D.F., México México, D.F., México Librado Ortiz-Ortiz Kenneth S. Warren Director, Instituto de Investigaciones Professor Biomédicas Department of Medicine Instituto de Investigaciones Biomédicas New York University School of UNAM Medicine México, D.F., México New York, New York THE EDITOR Roberto R. Kretschmer, M.D., is Professor of Immunology at the Faculty of Medicine, Universidad Nacional Autónoma de México (UNAM), and head of the Division of Immu- nology, Unidad de Investigación Biomédica, Instituto Mexicano del Seguro Social, Mexico City/Division of Experimental Medicine, Faculty of Medicine (UNAM). Dr. Kretschmer received his M.D. from the Faculty of Medicine, UNAM in 1963. He pursued his training in Pediatrics and Immunology at the University of Tubingen, West Germany and at Children's Hospital Medical Center, Harvard Medical School, Boston , Massachusetts (1963—1968). He became an Assistant Professor of Pediatrics at the Pritzker School of Medicine, University of Chicago, and Director of Pediatric Immunology at Michael Reese Hospital Medical Center, Chicago, IL (1974—1978), before returning to Mexico to his present position. Dr. Kretschmer is a member of the American Association of Immu- nologists, Infectious Diseases Society of America, Clinical Immunology Society, American Academy of Pediatrics and American Academy of Allergy and Clinical Immunology, Society of Pediatric Research, New York Academy cff Sciences, Sociedad Mexicana de Inmunología (past president), Academia Nacional de Medicina (México) and Academia Nacional de la Investigación Científica (México) among others. He has received the "Gabino Barreda" award for distinguished studies (UNAM) (1963) and the "Eduardo Liceaga" award for research (Academia Nacional de Medicina, México) (1986). Dr. Kretschmer has been the recipient of fellowships from the Mexican and West German governments and from the Panamerican Organization of States, as well as research grants from the National Council of Science and Technology (CONACYT, México). He has pub- lished over 90 specialized papers and contributed to 18 books. His current research interests are in immunology of amebiasis, chemotaxis, and the immunoinflammatory capacitie s of the newborn. REFERENCES 1. Brandt, H. and Pérez-Tamayo, R., Amibiasis, La Prensa Médica Mexicana, México, D.F., 1970. 2. Ravdin, J. I. and Guerrant, R. L., A review of the parasite cellular mechanisms involved in the path- ogenesis of amebiasis, Rev. Infect. Dis., 4, 1185, 1982. 3. Gitler, C. and Mirelman, D., Factors contributing to the pathogenesis behaviour of Entamoeba histolytica, Annu. Rev. Microbiol., 40, 237, 1986. 4. Martínez-Palomo, A., The Biology of Entamoeba histolytica, Research Studies Press, John Wiley & Sons, Chichester, 1982. 5. Martínez-Palomo, A., Amebiasis. Human Parasitic Diseases Series, Vol. 2, Ruitenberg, F. J. and Maclnnis, A. J., Eds., Elsevier, Amsterdam, 1986. 6. Ravdin, J. L, Amebiasis. Human Infection by Entamoeba histolytica, John Wiley & Sons, New York, 1988. 7. Warren, K. W., Foreword, in Amebiasis. Human Infection by Entamoeba histolytica, John Wiley & Sons, New York, 1988. 8. Walsh, J. A. and Warren, K. W., Selective primary care. An interim strategy for disease control in developing countries, N. Engl. J. Med., 301, 967, 1979. 9. Guerrant, R. L., The global problem of amebiasis: current status, research needs, and opportunities for progress. Introduction, current status and research questions, Rev. Infect. Dis., 8, 218, 1986. 10. Gallo, R. C. and Montaigner, L., The AIDS epidemic, in The Science of Aids, Piel, J., Ed., W. H. Freeman, New York, 1989, 1. 11. Petri, W. and Ravdin, J. I., Entamoeba histolytica, in Parasitic Infections in the Compromised Host, Walzer, P. D. and Genta, R. M., Eds., Marcel Dekker, New York, 1989, 385. 12. Schultz, M. G., Current concepts in parasitology, N. Engl. J. Med., 297, 1259, 1977. 13. W.H.O. Expert Committee on Malaria: sixteenth report, W.H.O. Tech. Rep. Ser., 549, 1, 1974. 14. Carpizo, J., Del Médico. Homenaje a Bernardo Sepúlveda, Universidad Nacional Autónoma de México, México, D.F., 1987.