VIROLOGY MONOGRAPHS DIE VIRUSFORSCHUNG IN EINZELDARSTELLUNGEN CONTINUING /FORTFÜHRUNG VON HANDBOOK OF VIRUS RESEARCH HANDBUCH DER VIRUSFORSCHUNG FOUNDED BY /BEGRÜNDET VON R.DOERR EDITED BY /HERAUSGEGEBEN VON S. GARD · C. IIALLAUER · K. F. MEYER 4 1968 SPRINGER-VERLAG WIEN GMBH THE INFLUENZA VIRUSES BY L. HOYLE 1968 SPRINGER-VERLAG WIEN GMBH ISBN 978-3-7091-2048-4 ISBN 978-3-7091-2046-0 (eBook) DOI 10.1007/978-3-7091-2046-0 All right-1:1 reservf'..d No part of this book mny be tranl'liated or reprodueed in any form witlwut written permission from Springer-Verlag © 1968 by Springer-Verlag Wien Library of Congre~s Catalog Card Number 68-54069 The Influenza Viruses By L. Hoyle Director, Public Health Laboratory, Northampton, England With 58 Figures Table ol Contents Preface 9 I. HistoricaJ Survey . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 l. Isolation of the Human Virus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 2. Neutralizatio!l and Complement Fixation . . . . . . . . . . . . . . . . . . . 10 3. The Haemagglutination Reaction . . . . . . . . . . . . . . . . . . . . . . . . . . 11 4. Protective Vaccination . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 5. Inhibitors of Haemagglutination . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 6. Intraccllular Reproduction of the Influenza Virus . . . . . . . . . . . . 12 7. Epidemiological Studies. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 8. Vaccination with Live Virus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 9. A.nimal Influenza . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 10. Chemotherapy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 11. Quantitative Relationships . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 II. Notnenclat.ure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 III. Classification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 L The ~fyxovirus Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 2, Sub-division of the Myxovirus Group . . . . . . . . . . . . . . . . . . . . . . 19 3. Classification of the Influenza Subgroup of the Myxoviruses. . . 19 IV. Isolation of the Influenza Viruses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 l. Source of Material . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 2. Isolation in Ferrets. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 3. Isolation in Mice . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 4. Isolation in Rat;; . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 5. Isolation in Hamster~ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.'5 6. Isolation in EggR . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 7. hmlat.ion in Tist~ue CuHnres . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 8. U ~e. of Virus Concentmtion Method~ in Isolation . . . . . . . . . . . . . 28 9. Relative Effieie,nr·y of Variou1; 1\Iethods of Virus Isolation . . . . . 28 Virol. Monogr. 4 2 L. Hoyle: The Influenza Viruses V. Cultivation of Influenza Viruses in Fertile Eggs . . . . . . . . . . . . . . . . . . . 28 l. Cultivation on the External Surface of the Chorioallantoic l\fembrane . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 2. Cultivation in the Amniotic Sac ........ 29 0 • 0 • • • • • • • • • • • • • • • • • ~l. Cultivation in the Embryo and Yolk Sa.c . . . . . . . . . . . . . . . . . . . . . 29 4. Cultivation in the Allantoic Sac . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 5. Cultivation in De-embryonated Eggs . . . . . . . . . . . . . . . . . . . . . . . 29 VI. Tissue Culture 11ethods for Influenza Virus Cultiva.tion. . . . . . . . . . . . . 30 1. Plaque )Iethods with Influenza Viruses . . . . . . . . . . . . . . . . . . . . . 32 2. Organ Cultures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 VII. Pmificat.ion of tbe Influenza Virus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 1. Purification by Centrifugation Methods . . . . . . . . . . . . . . . . . . . . . 33 2. Purification by Filtration Mct.hods .. , . . . . . . . . . . . . . . . . . . . . . . 34 3. Purification by Adsorption-Elution from Red Blood Cells . . . . . 34 4. Combined u~e of Adsorption-Elution from Red Cell~ and Dif- ferential Centrifugation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 5. Adsorption-Elution from Inhibitors . . . . . . . . . . . . . . . . . . . . . . . . 35 6. :-; ParLiek . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (j7 H. Corwlm;iow-< . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68 Table of Contents 3 XII. Resistance to Physical and Chemical Agents . . . . . . . . . . . . . . . . . . . . . . 68 l. Heat Resistance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69 2. Resistance to Freezing and Thawing. . . . . . . . . . . . . . . . . . . . . . . . 69 3. Re&istance to Drying . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70 4. Resistance to Ultrasonic Vibration . . . . . . . . . . . . . . . . . . . . . . . . 70 5. Resistance to pH Changes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70 6. Effect of Ions on Resistance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70 7. Resista.nce to Ionising Radia.tions . . . . . . . . . . . . . . . . . . . . . . . . . . 70 8. Resistance to Chemical Agents . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72 9. Survival and Resistance of Influenza Virus in Air . . . . . . . . . . . . 7 4 10. Inactivation of Yaccines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75 XUI. The Infective Property of the Influenza Viruses . . . . . . . . . . . . . . . . . . . 75 l. lnfectivity Titration in Mice. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77 2. lnfectivity Titration in Fertile Eggs . . . . . . . . . . . . . . . . . . . . . . . i7 3. lnfectivity Titration in Tissne Cultures . . . . . . . . . . . . . . . . . . . . . 78 4. Infectivity of Influenza Virus Ribonucleic Acid . . . . . . . . . . . . . . 79 5. Some Reflections on the Nature of the Infecting Property of Influenza Viruses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80 XIV. Haemagglutination . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81 l. The Salk Test . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81 2. Dcnsitometric Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82 3. Other .M:cthods of Haemagglutinin Titration . . . . . . . . . . . . . . . . 83 4. Effect of pH, Tempemture and Salt Goncentration on Haem- agglutination . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83 5. Mechanism of the IL!.emagglutination Reaction . . . . . . . . . . . . . . 84 6. Effcct of Virus on Lhe Red Cells . . . . . . . . . . . . . . . . . . . . . . . . . . . 86 7. Nature oE the Red Cell Receptors . . . . . . . . . . . . . . . . . . . . . . . . . . 87 8. Agglutination of Red Cells of Different Animal Species . . . . . . . 88 9. Relation bctween the Number of Virus Particles Present. in a ViruB Suspension and the Jla.emagglut.inin Titre. . . . . . . . . . . . . . 89 XV. Enzymic Activity of Influenza ViruReil . . . . . . . . . . . . . . . . . . . . . . . . . . . 89 l. Nature of the Virus Rceeptor . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90 2. 'l'he Reeeptor Destroying Enzyme of V. d10lerae, RDE . . . . . . . 91 3. Estimat.ion of Neuraminidase Aetivity . . . . . . . . . . . . . . . . . . . . . 91 4. J\.lechanism of Nemaminidase Action . . . . . . . . . . . . . . . . . . . . . . . 92 5. Purification of Viruil Neuraminirla~e . . . . . . . . . . . . . . . . . . . . . . . ()·) 6. Variations in t.he Neumminidase Aetiviiy of Different ::'tmins of Virus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93 7. Relation of Neurnminidasc to Haemagglut.inin . . . . . . . . . . . . . . 93 8. Relu,tion of Neuraminidase to t.lw Virus Growth Cyde . . . . . . . . 94 XVI. Inhibitors of Hacmagglutirmtion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95 l. 'l'he Frauci<> or ~ Inhibitor"... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9;) 2. The Chu or ß Inhibitor>' . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97 3. Ot.her Type~ of Ilamna,gglntiniu luhibit.or~ in Serum . . . . . . . . . !)8 4. Eledrophorcti<' Analyc;iH of Serum Inhihüors . . . . . . . . . . . . . . . . 98 5. Na,ture of the Non-speeifi<· Inhihit.on; of Normal Sera 119 4 L. Hoyle: The Influenza Viruses XVII. Serological Rea.ctions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100 A. Serologkai Reactions of the Inner Ribonueleoprotein Componeut .. 100 l. The Group Complement Fixation Test ..................... 100 2. Teclmique and e~e of the Group Complement Fixation Reaction 101 3. Precipitin Reactions with the SAntigen .................... 103 4. Specificity of the S Antigen ............................... 103 B. Serological Reactiou~ of thc Outer Virus Component ............ 104 l. The N eutralization Te8t . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104 2. Quantitative X eutralization Te~ts in Mice 104 :J. Quantitative X entralization rrest.s in Eggs .................. 105 4. Neutralization in Tissue Cultures .......................... 105 5. ~Iecha.nism of the X eutralization Reaction . . . . . . . . . . . . . . . . . . 106 6. The Haemagglutination-inhibition Test ..................... 107 i. Destruction of Xon·specific Inhibitors in Serum ............. 108 8. Antigenie Analysi,;; by the Use of I! I Tests ............. , .... 109 9. The Specific Cornplement FixfLtion Test .................... llO 10. Technique of the Specific Complement Fixation Test . . . . . . . . . llO 11, Agglutination, Pre<'ipitation and Flocculation Test'! .......... 111 12. Precipitin Reactiou~ by the Gel-diffusion Technique ......... 112 13. Antibody Abwrption 'l'eehuiques . . . . . . . . . . . . . . . . . . . . . . . . . . 112 14. Relation between Xcutralizing, Haemagglutinin-inhibiting, Speeific Cornplement l<'ixing and Preeipii.ating Antibodies .... 113 X\' III. Growth Cyele of the Influenza Virus - General Dcscription ......... 114 1. Sorrw Quantitative Considerations . . . . . . . . . . . . . . . . . . . . . . . . . 115 2. Intraeellulfl.r Reproduction of thc Influenza Viruses . . . . . . . . . . 116 XIX. Stage,.; in t.he Growth Cycle of Infhease to the Propertiell of ihe Virus 190 2. l'athology ot' Influenza ................................... l!H XX V I. .\letaholiflm of the lnfcded Cell in Influenza ...................... 197 XXVII. Lahoratory DiagnosiH of Influenza ............................... 201 A. Isolation and Identifieat.ion of Virus .......................... 201 B. ~erologiea1 DiagnoHis ....................................... 201 C. Rapid l\letl1ods of L;tboratory Diagnosis ...................... 204 XX VIII. lrnmunity anti Antibody Prod~etion in Influenza ................. 205 l. 1\atural lmrnunity to lnfluenr.al Infection .................. 205 2. Aequired lrnrnunit.y ...................................... 20G

The Influenza Viruses

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