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The Source and Action of Immunoglobulins in the Dog Intestine

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The Source and Action of Immunoglobulins in the Dog Intestine 2i .j.")-¡, THE SOURCE AND ACT]ON OF IMMUNOGLOBULINS IN THE DOG INTESTINE by Robent John Heddle, M.8., B.S ) Depantment of Micnobioì-ogy, Univensity of Adelaide Thesis submitted fon the degnee of Docton of Philosophy, Univensity of Adelaide Apni 1 19 76 ',t ,,'," :.1'. TABLE OF CONTENTS Page SUI"O4ARY I DECLARATION t-l-l ACKNOV,ILEDGEMENTS IV CHAPTER I INTRODUCTION I A The Impontance of Host-panasite Intenactions and of Othen Events at Mucous Membnanes in Continuity with the External Envinonment t B Evidence that Pnevious Expenience of a (Potential) Pathogen Can Prevent Disease Attnibutable to trl,ocalrt Infecti.on by that Pathogen 2 Development of the Concept of the Local- Immune System 4 7 The poo:r cornelation of senum and secnetory antibody titnes and of serum antibody titnes and local- immunity 4 2 The isol-ation of IgA and the demonstration of its abundance in a chanactenistic fonm in rrextennal secnetionstl 5 rrsecretony D The Present Concept of the IgA Systemrr 7 t Synthesis of IgA 7 2 The secretion of IgA 7t 3 Resistance to pnoteolysis of sec::etony IgA T2 4 Actiwities of secnetony IgA antibodies against micro-o:rganisms 16 5 Majon deficiencies in pnesent knowledge of the rrsecr.etony IgA system. rl 19 .t Roles of Antibodies of othen Classes and of Antibody Fnagments in Extennal Secnetions 22 7 (and 2.) Synthesis and secnetion 22 3 Susceptibility to pnoteolYsis 2B 4 Antibactenial activities 28 5 ltre antibactenial activities of antibody fnagments 30 Table of Contents (Cont. ) Page F Aims of the Study 30 I Oniginal aims 30 2 [Í¡dification of the oniginal aims 32 G Cholena as a Mode1 Disease 33 t The localised natune of the human and (selected) model diseases 33 rrantibacter- 2 The natur-e of ímmunity to cholena - ial-rt on rrantitoxicrr 34 3 The infant mouse cholena nodel 36 4 Penspectives on cholera as a model fon the nol-e of immunity in a localised bactenial infection of the gut 37 5 Cell mediated immunity and choler-a 38 H The rrsecnetony IgA Systemrr of the Dog 3B CHAPTER II MATERIALS AND METHODS 39 A. Anima-l.s 39 B. Sample Collection and Storage 40 I. Collection of samPles 40 2. Storage of samPles 43 C. Techniques fon Fnactionation of fmmunoglobulins 44 t. GeI filtnation 44 2, Ion exchange chnomatognaPhY 45 3. Affinity chnomatognaPhY 46 4. Concent::ation and dialysis of samples 4B D. fmmunochemical Techniques 49 t. rrOuchtenlonYtt analYsís 49 2. ImmunoelectnoPhonesis ( IE ) 50 3. Single nadial immunodiffusion ( SRID ) 51 4. Quantitative PreciPitins 52 Table of Contents (Cont. ) Page D Antisena 52 7 Antisena to r.¡hofe dog serum and whole dog colostrum 52 2 Refenence antrse:ra 53 3 Goat antisera specific fon dog -r þ on y chains - tt1\ccrr, ttAU tt and ttAYtr 54 F Measr:.r:ement of Pnotein Concentr:ations - Immuno- globulin Standa:¡ds 55 G Rabbit Antisenum to Punified Cholenagen 56 H Maintenance of V.choferae 5698 and Immunisation Pnocedunes 57 ï In vitro Assays fon Antibodies to I/.chol,erae 5698 58 t Pnepai:ation of V.choLerae 5698 lipopol.y- sacchanide (5698 LPS) 5B 2 Absonption of antj.bodies with 569B LPS followed by pnotein estimations 59 3 Passive haemaggJ-utination ( HA ) using sheep ned blood ce1ls sensitised with v.choTerae 5698 LPS (s6eB LPS HA) 60 4 The itCoombst rr test - enhancement of (passive) haemagglutinatíon with anti-dog heavy chain antisena 61 5 The "vibniocidal" assay - killing of v-choLetae 5698 by antibody and complement Ín vitro 61 6 Vibniocidal assays modified to assess the effects of lysozyme 63 7 Tneatment of antibodies with 2-mencaptoethanol 63 I Rernoval of antibody activity with v.cholerae 5698 LPS 64 trmaximum 9 Detenmination of means and possiblerr tib:es in senological assays 64 J In vívo Assays fon Antibodies to v.cholerae 5698 64 t The itopsonictt assay 64 2 The infant mouse cholena model 66 3 Removal of antibody activity with V.choTetae 569¡ LPS 68 Table of Contents (Cont. ) Page K. Studies on the Stability of Antibody Activities in Small Bowe1 Fluids 6B t In vitro studies 68 2 In vivo studies 70 3 Rabbit antibodies exposed to smalI bowel fluids 77 L rn vitro Pnocedures with Radio-iodine (131r ) 77 t Radio-iodínation using chlonamine-T 7t 2 Sepanation.of fnee and pr:otein bound nadio-iodine 72 3 Counting and the detenmination of enrolrs of count- ing nates 72 4 Radio-immunoelect::ophonesis and immuno-pnecipita- tion of radio-label 73 M In vivo Studies of the T::anspont of Dog Immunoglob- ulins fnom Senum into the fntestinal Lumen 75 7 Cneation of intestinal (Thiny-Vella) Ioops and cane of dogs 75 2 P::epanation of labelled immunoglobulins 76 3 Detenmination of inmunogJ-obulin üranspont 76 a I:r:rigating solution 76 b Basal sampling 77 c Administration of labeÌled ímmunoglobulins 77 d Penfusion of Thiny-Ve1la looPs 7B Autopsy of dogs and prepanation of tissue sections fon staining 79 f. Handling of penfusates and sena 79 g. Cytological studies of penfusates BO h. Assessment of samples using single nadial immunodiffusion (Snfl), nadio-assays and the vibniocidal assay 81 rtcleanancett itappeanance l_ Calculation of and rates tr 83 l Cornection for natunal tfvib:riocidalrl actívities 84 4 trPneliminany tt expeniments 85 5 Estimation of senum volumes and. of t,}.e in vivo half-life o¡ 131¡-¿oB IS2",b 85 Table of Contents (Cont. ) Page CHAPTER IIÏ STUDIES OF THE IMMUNOGLOBULTNS OF DOG EXTERNAL SECRETTONS 87 A fntnoduction 87 B Punification of Dog IgG 88 Punification of Dog IgM 90 D Punification of dog Secr.etony IgA 91 E Funthen Assessment of the IgM and Secnetony IgA Standands 92 F Immunochemical Studies of Dog Senum 94 G Immunochemical Studies of Dog Saliva 94 H Immunochenical Studies of Dog Mammary Secnetions 96 I Immunochemical Studies of Dog Sma1l Bowel Secnetions 9B t Snall bowel secnetions obtained at autopsy 98 2 SmalÌ bowel secr"etions obtained by penfusion in situ of acute loops of small bowel 100 CHAPTER IV THE ST]I'fi.,LATION AND PURTFICAT]ON OF SPECIFÏC ANTIBODIES T0 I/. choLerae 5698 WITH SPECïAL REFERENCE T0 lgA ANTïBODIES 105 A fntnoduction 105 B Estimation of Antibody to V.chol-erae 5698 Lipopoly- sacchanide (5698 tPS) 105 C Responses to Immunisation with T.cholerae 5698 113 t Panentenal immunisation 113 2 Loca1 immunisation of the panotid gland 116 3 Onal immunisation 721 4 Stimulation of antibodies in mammary secnetions t25 D Data, Denived fnom the Specific Activities of fmmunoglobu.l-ins in Secnetions and Senum, Relating to Local Synthesis and/on Tbanspont fuom Senum of Antibodies in Extennal Secretions 729 E Evidence that the Anti-v. choTerae 5698 Activity of Pr:::!fied Dog IgM on IgG Lay in the Immunochernically .Dominant Class 133 Table of Contents (Cont. ) Page F 'rVibniocidalil ancL Passive (Rnti*S0gB LPS) Haemagg- lutinating Activities of Secnetony IgA Pr:epanations at Diffe:rent Stages of Punification 134 CHAPTER V THE I'ANTIBACTERIALil ACTIVITIES 0F DOG IgG, IgM AND SECRETORY IgA ANTIBODIES T0 y.chof,erae 569B 144 A fntnoduction 144 B rn vitro killing of V.cholerae 5698 with Complement - the Vibriocidal Assay !47 T The activities of the diffenent cl-asses of antibody 747 2 'rVibniocidalr activities of antibodies aften tneatment with mencaptoethanol 152 3 The nole of lysozyme in the víbniocidal neaction 154 C Opsonisation of V.chol-erae 569B 159 D. Pnotectíon of 5 - 6 Day Old Mice against 0ra1 Challenge with Y.cho-l.erae 5698 163 t Efficiencies of diffenent classes of antibody 163 2 The specificíty of the p:rotective antibodies t67 3 The possible nature of the antibactenial effect of p:rotective antibodies 168 CHAPTER VI THE EFFECTS OF DOG AND RABBIT SMALL BO}TEL FLUIDS ON FUNCT]ONAL PROPERTÏES OF DOG AND RABBIT ANTI- V. choTerae 5698 ANTIBODIES 775 A Intnoduction 775 B Pneliminany Obsenvations 776 t Single nadial immunodiffusion analyses in the pnesence of pnoteolytic enzymes 776 2 frnmunochemical studies of snall bowel washings t76 c. Effects of Smal1 Bowel Fluids on the I'Vibniocidalr Capacíties of Antibodies to T.cholerae 5698 179 t In vitro exposune of antibodies to small- bowel fluids (SBF) t79 a. Incubation of dog and :rabbit antibodies in homol-ogous sma1l bowel fluids 179 b. Effects of sma11 bowel fl-uids on vibniocidal assays 185 Table of Contents (Cont. ) Page c Kinetics of the l.oss of 'rvibniocidal-'t activity 187 d Incubation of dog IgG antibodies in nabbit smal-l bowel fluids 188 e. Confinmation of the pnoteolytic activities of dog and rabbit small bowel ffuids using t3t. --*iodine-labelled bovine serum albumin 1BB 2 In vivo exposune of antibodies to small bowel fluids 190 D Pnotective Activities of Dog Ig G2arb Anti-y. cholerae 5698 Antíbody rrDigests" 792 ! Punpose of the expeniments 792 2 Pr:otective and trvibniocidaltr activities of dog IgG antibody I'digeststt 792 3 The punity and specificity of the pnotective antibody 195 a. Immunochemical studies 195 b. Specificity of the pnotective antibody 195 c. Class of the pnotective antibody 196 131ï-Igcr.,o 4 Ge1 fil-tnation of '?digests'r of 196 CHAPTER VTI THE TRANSPORT OF DOG IMMUNOGLOBULINS FROM SERUM INTO THE LUMEN OF THE DOG SMALL INTESTTNE 200 A fntnoduction 200 7 Or"igins and tnanspont of intestinal IgA 200 2 Onigins and tnanspont of intestínal IgM and IgG 200 a. Significanee 200 b. P::evious studies 201 c. Expenimental appnoach 203 B. The Sungical Model 20+ 7 Anatomical, histological and functional featunes of intestinal (Thiny-Vella) loops 20+ a.
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