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• JL*J*• * •T * NN08201.B95 -fes- A QUALITATIVE AND QUANTlf*faf$ INVESTIGATION OF ,0L£ACtt).ItY*>\]fjPL • N*4S4L^l^SPIRA^ #RY MUCOSAL* j£ | % •t * SUHF»^IE»6#*€OW 4ND SH£EP « II m mm i m • #ji It * li « < m m W 41 9 it 11 in • JL*J*• * • * wI « • • NN08201.B95 636.2+ 636.4:591.872:591.185.34:591.487 B. P. M. MENCO A QUALITATIVE AND QUANTITATIVE INVESTIGATION OF OLFACTORY AND NASAL RESPIRATORY MUCOSAL SURFACES OF COW AND SHEEP BASED ON VARIOUS ULTRASTRUCTURAL AND BIOCHEMICAL METHODS (with a summary in Dutch) PROEFSCHRIFT TER VERKRIJGING VAN DE GRAAD VAN DOCTOR IN DE LANDBOUWWETENSCHAPPEN, OP GEZAG VAN DE RECTOR MAGNIFICUS DR. H. C. VAN DER PLAS, HOOGLERAAR IN DE ORGANISCHE SCHEIKUNDE IN HET OPENBAAR TE VERDEDIGEN OP WOENSDAG 12 OKTOBER 1977 DES NAMIDDAGS TE VIER UUR IN DE AULA VAN DE LANDBOUWHOGESCHOOL TEWAGENINGEN H. VEENM AN & ZONEN B.V. - WAGENINGEN - 1977 STELLINGEN I Vries-ets resultaten tonen aanda t olfaktorische cilia zich in hun partikel- populatie onderscheiden van andere nasale haartjes. MENCO, B.P . M., 1977. Freeze-etch morphology of olfactory and respiratory nasal hairlets inrat ,co w andfrog . - In: Olfac­ tion and Taste,VI. :I n Press. II Daar debewegin g vancili a ookzonde r omringend membraan geschiedt is het onwaarschijnlijk dat vries-ets partikels, gevonden in de intramembraan breukvlakken van olfaktorische cilia, een substantie vertegenwoordigen welke perse noodzakelijk isvoo r cilium beweging. Dit proefschrift. WIEDERHOLT, NJ- L., 1976.Mechanosensor y transduction in 'sensory' and 'motile' cilia. - Ann. Rev.Biophys . Bioeng., 5: 39-62. Ill Van cilia afgeleide sensorische receptor structuren zijn in te delen in twee hoofdgroepen; ten eerste een hoofdgroep waarbij het receptor mechanisme berust opmechanisch e verstoringen inhe treceptormembraa n ente n tweede een hoofdgroep waarbij hetreceptormembraa n specifieke receptorstrukturen bevat welkeinterakti eme t destimulu s aangaan. Dit proefschrift. BARBER, V.C , 1974.Cili ai n senseorgans .- In: Ciliaan d Flagel- la, SLEIGH, M.A. , ed.: 403-433,Acad . Press, London. MORAN, D.T., ROWLEY III, J. C, ZILL, S. N. & VARELA, F. G., 1976. The mechanism ofsensor y transduction ina mechanoreceptor. Functional stages in campaniform sensilla during the molting cycle,J . Cell Biol., 71: 832-847. IV In tegenstelling totd eopvattin g van Getchell enGetchel l zijn ermee r over- eenkomsten tussen geurstof-receptor interaktie en hormoon-receptor inter­ aktie dan tussen geurstof-receptor interaktie en substraat-enzym interaktie. DODD, G. H., 1976. Ligand-binding phenomena in chemo- reception. - In: Structure-Activity Relationships in Chemo- reception, BENZ, G., ed.: 55-63, Information Retrieval Ltd., London. GETCHELL, T. V. & GETCHELL, M. L., 1977. Early events invertebrat e olfaction. - Chem. Sens. Flav., 2: 313-326. V Hetverdien t aanbevelingd eexponentiel e funktie, zoalsgebruik t wordt ind e psychofysika, voor geur- ensmaakonderzoe k tevervange n door eenasymp - totische. HECK, G. L.& ERICKSON,R . P., 1973. Arat e theory of gustatory stimulation. - Behav. Biol., 8: 687-712. PUNTER, P.H. , 1976. Odor intensity for n-butanol; individual vs. groupdata .- Proc.2n dECR O Congress, Reading, England: 37. VI Deorganoleptisch e kwaliteit vannatuurlijk e extracten kan vertroebeld wor- dendoo r bijprodukten dietijden s de 'processing' van het voedingsmiddel ont- staan. TASKINEN, J. & NYKANEN, L., 1976. Volatile constituents of an alcoholic extract of juniper berry. - Flavours, September/ October 1976: 228-229. VII De bewering van Meritt en medewerkers dat genetische varianten vana - amylase uit menselijk speeksel electroforetisch langzamer zouden zijn dat het meest voorkomende isozym-patroon, berust ophe t feit datsneller e varianten bijheterozygote nme td edoo rhe ngebruikt etechnieke nnie twaarneembaa rzijn . MERRITT, A. D., RIVAS, M. L., BIXLER, D. & NEWELL, R., 1973. Salivary andpancreati c amylase: Electrophoretic characteriza­ tions and genetic studies.- Am.J .Hum . Genet., 25:510-522 . PRONK, J., 1977.A genetic variant of amylase from human parotid saliva detected by isoelectric focusing. - In: Electro- focusing and Isotachophoresis. RADOLA, B. J. & GRAESSLIN, D., eds.: 359-366,Walte r de Gruyter &Co. , Berlin. VIII Het feit dat 'De Emmausgangers' van Han van Meegeren in Museum Boymans-Van Beuningen geen naambordje draagt, zou bij dehuidig e direktie eenzelfde hoeveelheid schaamrood opd e wangen moeten brengen als bijd e direktiedi eindertij d tot aankoop isovergegaan . B. P. M. MENCO Wageningen, 12oktober197 7 'This country', said Eliot, 'had tremendous research projects devoted to fighting odors. They were supported by individual contributions given to mothers whomarche d on Sunday from door todoor . The ideal of the research was to find a specific chemical deodorant for every odor. But then the hero, whowa sals oth ecountry' sdictator ,mad ea wonderfu l scientific breakthrough, even though he wasn't a scientist, and they didn't need the projects anymore. He went right to the root of the problem'. 'Uh huh', said the Senator. He couldn't stand stories by Kilgore Trout, and was embarrassed for his son. 'He found one chemical that would eliminate all odors?' he suggested, to hasten the tale to a conclusion. 'No. AsI say,th eher owa sdictator , and hesimpl yeliminate d noses.' From: 'God BlessYou , Mr. Rosewater' by KURT VONNEGUT JR. Tomy Parents, Idoand Marion, and everybody who was involvedin preparing this thesis CONTENTS GENERAL INTRODUCTION 1 1. MACROSCOPIC AND MICROSCOPIC STUDIES ON BOVINE NASAL EPI­ THELIUM USING LIGHT MICROSCOPY, LOW- AND HIGH-VOLTAGE TRANSMISSION AND SCANNING ELECTRON MICROSCOPY 3 1.1. Introduction 3 1.2. Materials and methods 4 1.2.1. Methods identical for all microscopic investigations 4 1.2.2. Preparation of sections for high-voltage transmission observation ... 5 1.2.3. Preparation of ultrathin sections for normal transmission electron micro­ scope observation 6 1.2.4. Preparation of sections for light microscopic observation 6 1.2.5. Preparations oftissu eblock s for examination ina scannin g electron micro­ scope 6 1.2.6. Account of the amount of tissue material used 7 1.2.7. Materials 7 1.3. Results 7 1.3.1. Macroscopic appearance 7 1.3.2. General histology of the bovine nasal mucosa 8 1.3.3. Structure of the bovine respiratory epithelium 10 1.3.3.1. Scanning electron microscopy of the bovine respiratory epithelium ... 10 1.3.3.2. Transmission electron microscopy of the bovine respiratory epithelium . 11 1.3.4. Structure of the bovine olfactory epithelium 15 1.3.4.1. Scanning electron microscopy of the bovine olfactory epithelium .... 15 1.3.4.1.1. General anatomy 15 1.3.4.1.2. Temporal and spatial differences in the surface of the bovine olfactory epi­ thelium as shown by the scanning electron microscope 25 1.3.4.2. High-voltage and normal transmission electron microscopy of the bovine olfactory mucosa 25 1.3.4.2.1. General description of the mucosa 25 1.3.4.2.2. A description of the non-ciliary part of the olfactory mucosa 27 1.3.4.2.3. Temporal and spatial differences in the bovine olfactory epithelium as seen by transmission methods 30 1.3.4.2.4. Ciliary and microvillous structures 35 1.4. Discussion 46 1.4.1. General 46 1.4.2. Expansion and turn-over of the olfactory epithelium 47 1.4.3. Nasal epithelium surface structures 47 1.4.3.1. Endocytotic vesicles within the olfactory nerve ending 47 1.4.3.2. Brush cell 49 1.4.3.4. Ciliary structures 50 1.4.3.4.1. General 50 1.4.3.4.2. Axonemal aggregates 50 1.4.3.4.3. Basal bodies 50 1.4.3.4.4. Ciliary axonemal and membrane structures 51 1.4.3.5. Electron-lucent mucus inclusions 54 2. A FREEZE-ETCH AND ELECTRON SPIN RESONANCE STUDY ON NASAL EPITHELIUM OF COW AND SHEEP 56 2.1. Introduction . 56 2.2. Materials and methods 57 2.2.1. Freeze-etch studies 57 2.2.2. Electron spin resonance studies 57 2.2.3. Materials 59 2.3. Results 59 2.3.1. Freeze-etching 59 2.3.2. Electron spin resonance studies on sheep olfactory and respiratory nasal mucosa . 63 2.3.2.1. Order parameters 63 2.3.2.2. Label incorporation ratios 72 2.3.3. ESR studies on odorant interacted model membranes 78 2.4. Discussion 78 2.4.1. Junctional complexes as seen by the freeze-etch technique 78 2.4.2. Axons 79 2.4.3. Microvilli 80 2.4.4. Ciliary necklaces 80 2.4.5. Ciliary fracture faces above the necklace 80 2.4.6. Results of freeze-etch and spin label experiments in relation to other re­ ceptor systems 82 3. QUANTITATIVE ANALYSES OF CILIATED AND MICROVILLI BEARING SURFACE STRUCTURES OF THE BOVINE OLFACTORY AND NASAL RESPIRATORY EPITHELIUM 85 3.1. Introduction 85 3.2. Materials and methods 85 3.3. Results ' 87 3.3.1. Evaluation of measurements on some nasal structures. 87 3.3.2. Some parameters not included in the comparative evaluations 88 3.3.2.1. General cellular appearances of bovine respiratory and olfactory epithelia 88 3.3.2.2. Ciliary microtubule (axoneme) and membrane dimensions 88 3.3.3. Effect of the observation method on some dimensions and frequencies of epithelium structures of bovine olfactory and respiratory mucosae ... 88 3.3.4. Olfactory versus respiratory cilia 90 3.3.5. Effect of age on olfactory and respiratory epithelium structures .... 93 3.3.6. Comparison between olfactory and respiratory epithelia 93 3.3.7. Comparison of various olfactory regions 93 3.3.8. Number of cilia per nerve ending 94 3.3.9. Relationships between the observed structural parameters 94 3.3.10. Revised determinations of nerve ending densities 102 3.3.11. Volumes and surface areas of nerve ending structures and particle fre­ quencies 104 3.4. Discussion 106 3.4.1.
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