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The Physiology and Immunology of the Endocrine Testis

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The Physiology and Immunology of the Endocrine Testis tì/ jì;:llilul s\ t: 1.q_ 81 LI Àt{Y THE PHYSIOLOGY AND IMMUNOLOGY OF THE ENDOCRINE TESTIS by Simon MADDOCKS, B.Ag.Scl. (Hons) A thesls submltted to the Unlversfty of Adelalde ln fulfl'lment of the requlrements for the degree of Doctor of PhllosoPhY Department of Anlmal Scfences Walte Agrlcultural Research Instltute The UnlversitY of Adelalde March 1987 lì,.,ì,r(,:,td r/ riÈ"1 Know'ledge and wlsdom far f rom belng oner. Have ofl tlmes no connexion. Knorledge dwells In heads replete with thoughts of other men; Wlsdom ln minds attentlve to thelr otvn' Know'ledger a rude unprof ltab'le massr The meré materials with whlch wlsdom bul'ldsr Tfl.|smoothldandsquarldandflttedtoftsplacer Does but encumber whom lt seems trenrlch' Knowledge ls proud that he has learnrd so much; Wisdom ls humb'le that he knows no more' I{il'l lam CowPer ( 1731-1800) '_ .å Cowper Poetical Works TABLE OF CONTENTS f CONTENTS vii ABSTRACT x DECLARATION xl ACKNOt{LEDGEMENTS PREFACE xlff PART I: T CTIAPTER I: INTRODUCTION 2 1. 1. INTRODUCTION 2 L.2, THE PHYSIOLOGY OF THE ENDOCRINE TESTIS L.2,L. Introducti on 2 L.2.2. The Leydfg cells 7 r.2,2. I, Anatomy 7 L.2.2. 2. Relatiónship to blood vessels, lymph vessels and seminiferous tubul es L2 .2.2 3, Re]atfonshlp to other ce]ls in fnterstlt'lal tissue I5 I 18 ,212 4. Hormones produced by the Leydfg ce1ìs I 22 L .2.3 The Sertoll cells 24 T .2,3 I. AnatomY I .2.3 2. Relatlônshlp to the blood-testis barrier 29 I ,2,3 3. RelationshiP to other ce'lls 32 I 2.3 4. Hormones produced by the Serto'li cells 35 I .2.4 Mechanfs|n of transport of hormones out of and across cel'ls 38 L.2,4 .I.. Hormone secreti on 38 1.2,4.2. TransPort 4L L,2.5 . Cotpositìon and functlon of interstitial f'luid 45 L.2.6 . Conirol of endocrine activity in the testis 52 57 ,3 THE IMMUNOLOGY OF THE ENDOCRINE TESTIS I 57 T .3 Introducti on .r 58 ,3 r2 The Immune sYstem I 69 13 .3 Immunologica'lly privlleged sltes I 7L ,3 r4 The testis as án.fmmunologfcally privileged sÍte t 78 ..3 .5 TheorlesforthePrlvilegedstatusofthetestis I 84 I .4 THE PURPOSE OF THIS STUDY BART II: 86 CTIAPTER 2: MATERIALS AND METHODS 87 2.L. REAGENTS 87 2.2. ANIMALS 88 2. 3 SURGICAL PROCEDURES 88 2, 3 r f . ThyroidectomY 9I 2' 3 t2.. Hypophysectomy 93 2. 3 13r Cryptorch I dectomY 93 2. 3 Efferent Duct Llgatlon .4, 94 2. 3 .5. Blood col lectlon if 2.3.5 Rats 94 .I. 95 2.3.5 SheeP .2. 97 2.3.6 . Lymph col lectlon 2.3.7 . Rat TransPl ants 97 2.3.7 .I. Under the kfdneY caPsule 97 2.3.7 Into the testfs 98 .2. 99 2.3.8 . Sheep Transplants lnto the testis 99 2.4. ISOLATION AND CULTURE OF RAT PITUITARY CELLS 2.4.L. Buffers and SolutÍons 99 'l IOI 2.4.2. Co1 agenase d f gestl on 2.4.3. Trypsfn dlgestlon t02 2.4.4. Tlssue culture 103 Routine method 103 2.4.4. I. r03 2.4.4. 2. LHRH chal'lenge TESTS 104 2.5 . THYROID FUNCTION I04 Iod I ne accumu'l atf on 2.5 . 1. 104 2.5 . Rats l. I. 106 2.5 SheeP .L.2. 106 2.5 .2. TRH chal'lenge 108 2.6 . PITUITARY FUNCTION TEST - LHRH CHALLENGE 2,6 .I. Intravenous iniectlon 108 Intravenous lnfusion I08 2.6 .2. r08 2.6 -.2.2,,2,L. Va]idation of method Calculatfon of dose of fnfused LHRH r09 2.6 r09 2.6 .2.3. Intratesticular arterlal lnfuslon 110 2.7. HCG CTIALLENGE rl0 2.8. EDS ADMINISTRATION 1r0 2.9. SILICA ADMINISTRATION VOLUMES TIO 2.L0. MEASUREMENT OF ALBUMIN AND CT-EDTA DISTRIBUTION ttl z.LL. VASCULAR PERMEABILITY LL2 2.L2. LYMPH FLOW 113 2.T3. HORMONE ASSAYS Buffers and Solutions 113 2,L3,L. 113 2.L3.L,L. PhosPhate buffer L25 Blocker so]ution wlth I-thy roxi ne t14 2.L3.L.2. 114 2.L3 .L.3. TO-f ree serum 1r4 2.L3 .I.4. LÍquid scintfl'latlon system L-thYroxine radlofmmunoassaY T$ 2,r3 ,2. r16 2.r3 116 2,r3 L2l 2.L3 L23 2.L3 L24 2,L3 L25 2,L3 L27 2.13 Rat Prolactin (rPRL) .3.6. (oTSH) t28 2.L3 .3.7 . Ovlne Thyroid stlmul ating hormone ftf r29 2.L4. HISTOLOGY 2.I4.I. Ff xatlves r29 2.L4.2. Tfssue Processlng L29 13I 2.L5 . IMIVIIJNOH I STOCTIEMISTRY 2.L5.L. Flxatlves 131 2.L5.2. Tlssue Processlng 133 2.L6. STATISTICAL ANALYSIS 135 GTIAPTER 3: THE RODENT TESTIS AS AN IMMUNOLOGICALLY PRTVILEGED SITE 136 L37 3.1. INTRODUCTION r37 3.2. CHOICE OF TISSUE GRAFTS 3.3, PITUITARY TRANSPLANTS 138 3..3. I, Experlmenta'l Procedures 139 139 3.3, I . l. Anlmal s 3.3. I.2. Col I agenase-df spersed cel'l s 139 3,3. I.3. TrYpsin-disPersed cel'ls r40 r40 3 .3. I.4. Ouarters of whole Pltultarfes 141 3.3, 2 . Results 3.3. 2 Col1 agenase-df spersed cell s 14t .I. r42 3 2 ,2. Trypsln-dlsPersed cel'ls '3. 145 3.3. 2 .3. Ouarters of who'le pituitaries 3.3.3. Discusslon 150 il 156 ip 3 .4. THYROID TRANSPI-ANTS tI Experf menta'l 157 ;l 3.4.1 . Procedure 3.4.r .1. Anfma'ls r57 3.4.1 .2. Survival of thyroid allografts in the testis rs7 3 .4.L .3. A comparlson of graft survlval ln the kidney and the testis 158 3.4.2. Resul ts 158 3.4.2.r. Survival of thyroid allografts in the testis 158 3 .4.2.2. A comparison of graft survfval fn the kldney and the testls 159 3.4.3 Dfscussion 165 3.5. EXPERIMENTAL MANIPULATIONS OF THE TESTIS AND THEIR EFFECTS ON ALLOGRAFT SURVIVAL r69 3.5. I. Efferent Duct Ligation 169 3.5. 1. I, Experimental Procedure 17I 3,5. I. 2. Results 17I 3.5, t.3. Dlscusslon r72 L72 3.5. 2' Cryptorch Íd i sm 3.5. 2. l. Experimenta'l Procedure 175 175 3 .5, 2.2, Results 3.5. 2.3 . Dlscusslon L76 3.5, 3. Human Chorionic GonadotroPhin L76 3.5. 3.1 . Mafntenance of lncreased lymph flow wlth hCG L77 3.5.3,r ,1, ExPerfmental Procedure L77 3.5.3.r .2. Resu'lts 178 3.5.3.r .3. Dfscusslon r80 l lv 3.5.3.2. Effect of lncreased lymph f'lorv on allograft surv lval 184 r84 2.L. Experf mental Procedure 3.5.2. r85 3.5.3. 2.2. Resul ts 187 3.5.3. 2.3. Df scusslon 3.6. IMPoRTANcEoFLEYDIGCELLSAND/oRMACRoPHAGESFoR 188 ALLOGRAFT SURVIVAL IN THE TESTIS r89 3.6.1. Experlmental Procedure 190 3.6.2. Resul ts 3.6.3. Discusslon 195 CHAPTER4:THEOVINETESTISASANIMMUNOLOGICALLY 200 PRIVILEGED SITE 20L 4.I. INTRODUCTION 202 4.2. EXPERIMENTAL PROCEDURE 202 4.2.L. Animal s 4.2.2. Thyrold transPl ants 203 4.2,3 . Pitultary transPl ants 204 206 4.2.4. Immunohl stochenl strY 206 4.2.5. Hormone assays 206 4,3, RESULTS 4.3. r. Thyrold transPl ants 206 206 l_ TRH challenge 4.3. r. 208 4 2 Iodine local lsatlon 13 .L. 208 4..3. 3 Hi sto'logY 'j r. 208 fr,| 4 .3 4 Plasma ãn¿ lymph hormone concentrations ì{ ,1. 2L2 4,.3.1. 5 P]asmahormoneconcentratÍonsfnweek]ysamples È 2L2 4.3 .2, Pitultary transPl ants 2L2 4.3 LHRH challenge ,2, L 2L2 4.3 .2, 2. Plasma and lymph hormone concentrations 215 4.3,2. 3. Immunohl stocheml strY 4.3.2. 4. plasrna hormone concèntrations in weekly samp'les 2t5 2r5 4.4. DISCUSSIoN 2L8 DTSCUSSION PART III: 225 CTIAPTER 6: MATERIALS AND METHODS 226 6.1. REAGENTS 226 6.2. HEATING OF TESTES 226 6.3. PREGNENOLONE ADMINISTRATION t 227 I 6.4. TESTOSTERONE IMPLANTS I ¡ 227 6.5. CELL DISPERSION TECI{NIOUE ì v 228 ASSAY 6.6. HORMONE 228 6.6.1 Testosterone radi oimmunoassay 230 6.7. PROTEIN ASSAY reagent BL 6.7 .Lr Proteln 231 6.7.2. Proteln assay LEVELS B4 6.8. MEASUREMENT OF SODIUM AND POTASSIUM STUDIES 234 6.9. ADMINISTRATION OF RADIOISOTOPES IN PHYSIOLOGICAL 6.9.1. 3H-v,lut". 234 6.9.2. 3H-Testosterone 235 14c-Mann 235 6.9.3 . ltol 125r-No.*u] 236 6.9.4. Rat Pl asma 6.9.5. 22Nu-sodium Chloride 236 237 6.T0. STATISTICAL ANALYSIS 238 REGION 239 7.L. INTRODUCTION 24L CANNULA 7 .2, THE PUSH-PULL 24L on 7.2,L, Introductl 242 7.2.2' Construction 245 Apparatus 7 ,2.3, 249 7.2.4. Usä of the Push-Pu11 cannula 249 IN THE RAT 7 .3. INVESTIGATIONS 25r 7.3.1. Experimental Procedure 'App'l in the testis 25L 7 .3.L, I. ication 25L 7 ,3.L, 2, Measurement of dilution Chofce of infusates 253 7 ,3 ,1, 3. 253 7 4. Protocol used .3 'Lr 254 7 ,3,2. Resu'lts Applicatfon ln the testls 254 7 .3.2, I. 254 7 .3.2' 2. Measurement of dllutlon of interstitfal flufd 258 7 .3.2.3. AnalYsls 258 7 .3.3.
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