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I. Gametogenesis ASPECTS OF GAMETOGENESIS AND RADIATION PATHOLOGY IN THE ONION FLY, HYLEMYA ANTIQUA (MEIGEN). I. GAMETOGENESIS J.A.B.M.THEUNISSEN ASPECTS OF GAMETOGENESIS AND RADIATION PATHOLOGY IN THE ONION FLY, HYLEMYA ANTIQUA (MEIGEN). I. GAMETOGENESIS Dit proefschrift met stellingen van JOHANNES ANTONIUS BERNARDUS MARIA THEUNISSEN landbouwkundig ingenieur, geboren te 's-Gravenhage op 10 april 1937, is goedgekeurd door depromotoren , Dr. J. deWilde , hoogleraar inhe t dierkundig deelva n de Plantenziektenkunde en mej. Dr. L. P. M.Timmermans , lector in dealgemen e dierkunde De Rector Magnificusvan de Landbouwhogeschool, J. P. H. VAN DER WANT Wageningen, 12 april 1976 This thesis is also published as Mededelingen Landbouwhogeschool Wageningen 76-3 (1976) (Communications Agricultural University Wageningen, The Netherlands) 632.773.4 595773.4:591.46 J. A. B. M. THEUNISSEN ASPECTS OF GAMETOGENESIS AND RADIATION PATHOLOGY IN THE ONION FLY, HYLEMYA ANTIQUA (MEIGEN). I. GAMETOGENESIS (with asummary inDutch) PROEFSCHRIFT TER VERKRIJGING VAN DE GRAAD VAN DOCTOR IN DE LANDBOUWWETENSCHAPPEN, OP GEZAG VAN DE RECTOR MAGNIFICUS, DR. IR. J. P. H. VAN DER WANT, HOOGLERAAR IN DE VIROLOGIE, IN HET OPENBAAR TE VERDEDIGEN OP VRIJDAG 21 MEI1976 DES NAMIDDAGS TE VIER UUR IN DE AULA VAN DE LANDBOUWHOGESCHOOL TE WAGENINGEN H. VEENMAN&ZONEN B.V.-WAGENINGEN-1976 TAB: XL II. ru.a. r#.v. rjf.i. TyJB. Male (fig. VII) and female (fig. VIII) reproductive organs of Tabanus, as has been drawn by the Dutch Entomologist Jan Swammerdam around 1669. Source: Joannis Swammerdammie, Biblia Naturae sive Historia Insectorum, Leyden MDCCXXXVIII. (University Library, Wageningen). STELLINGEN De definities van de begrippen 'predefinitive', 'indefinitive' en 'definitive' spermatogonien zoalsz e worden geformuleerd door Hannah-Alava, leidento t de conclusieda t decategori e van de'indefinitive ' spermatogonien overbodigis . Dit proefschrift. HANNAH-ALAVA, A. 1965. Adv.i n Genetics, 13, 157-226. II De invoering vand eter m 'ovarioles adenomorphes' door Martoja ter aan- duiding vanhe t ovariooltype van Steraspisspeciosa isoverbodi g enongewenst . MARTOJA, R. 1964.Bull . Soc. Zool. Fr.,89 ,614-641 . HI De conclusie vanCausse , Feron en Pereau-Leroy dat in Ceratitis capitata het verdwijnen van de spermatiden binnen vier dagen nabestralin g moet wor­ den toegeschreven aan voortgezette differentiatie, wordt ten onrechte ge- trokken. CAUSSE, R., FERON, M. en PEREAU-LEROY, P. 1968. in: "Radia­ tion, radioisotopes and rearing methods in thecontro l of insect pests", 355-363, International Atomic Energy Agency, Wenen. IV Beoordeeld naar histopathologische criteria zijn germinale cellen bij de uievlieg niet altijd gevoeliger voor ioniserende straling dansomatisch e cellen. Dit in tegenstelling totd estrekkin g vand e'wet ' vanBergoni e enTribondeau , die debasi si sva n de radiotherapie. BERGONIE, J. en TRIBONDEAU, L. 1906. C. r. Acad. Sci., Paris, 143, 983, Onderzoek naar criteria voor 'kwaliteit' van gekweekte en/of behandelde insecten voor gebruik ingenetisch e bestrijdingsmethoden dient eenhog e prio- riteit te krijgen. CHAMBERS, D. L. 1975. in: "Controlling fruitflies b yth e sterile- insect technique", 19-32, International Atomic Energy Agency, Wenen. VI Het aantal bladluizen vand esoor t Myzus persicae SULZ. in pootaardappe- leni s geencorrec t criterium voor het doenbeeindige n van de groeiperiodeva n dit gewas. J. A. B. M. THEUNISSEN Wageningen, 21 Mei, 1976. VII Niet alleen insecten kunnen door Baculovirussen worden gei'nfecteerd. REED, D. K. en HALL, I. M. 1972. J. Invertebr. Pathol., 20, 272-278. VIII Stoffen die nematoden activeren tot wortelpenetratie, zoals 'hatching agents', behoeven niet noodzakelijkerwijze afgescheiden te worden in wortel- exudaten. CARROLL. K. K. 1958. Nematologica 3, 197-204. IX De dalingva n het aantal zeehonden ind eWaddenze e kan niet worden toe- geschreven aan deopnam e van kwik via voedselketens. Het vanuit onze samenlevingwille n bepalen welke graad van technologische ontwikkeling voor ontwikkelingslanden passend zou zijn, getuigt van elitair denken en een paternalistische houding tenopzicht e van deze landen. XI Alsd ezwijgend e meerderheid gaat spreken, leert desprekend e minderheid zwijgen. CONTENTS 1. INTRODUCTION 1 2. MATERIAL AND METHODS 2 2.1. Insect Material 2 2.1.1. Bionomics of the onion fly 2 2.1.2. Other insect species 5 2.2. Methods 5 2.2.1. Cytological methods 5 2.2.2. Phase contrast microscopic methods 5 2.2.3. Histological methods 6 2.2.4. Autoradiographic methods 8 2.2.5. Electronmicroscopic methods 9 2.2.6. Illustrations 10 2.2.7. Sampling 10 3. GAMETOGENESIS 11 3.1. Introduction 11 3.2. Gross anatomy of the reproductive organs 11 3.3. Cytological observations 16 3.4. Early post-embryonic development 18 3.5. Spermatogenesis 19 3.5.1. Introduction 19 3.5.2. Identification of testicular cell types 20 3.5.2.1. Structural organization of the young adult testis 20 3.5.2.2. Germinal cell types ; . 21 3.5.2.2.1. Primary spermatogonia 23 3.5.2.2.2. Secondary spermatogonia 33 3.5.2.2.3. Primary spermatocytes 38 3.5.2.2.4. Secondary spermatocytes 43 3.5.2.2.5. Early spermatids 45 3.5.2.2.6. Intermediate spermatids 49 3.5.2.2.7. Transformation spermatids 51 3.5.2.2.8. Sperm cells 57 3.5.2.2.9. General discussion on germinal cell types 57 3.5.2.3. Somatic cell types 64 3.5.2.3.1. Testicular sheath 65 3.5.2.3.2. Apical cell 70 3.5.2.3.3. Cyst cells 72 3.5.2.3.4. Central cavity cells 75 2.5.2.3.5. Basal cells 76 3.5.2.3.6. Terminal epithelium 76 3.5.2.3.7. Membrana basilaris 78 3.5.2.3.8. General discussion on somatic cell types 79 3.5.3. Testicular development 81 3.5.3.1. Larval development 81 3.5.3.2. Pupal development 84 3.5.3.3. Adult development 90 3.5.4. Dynamics of spermatogenesis 97 3.5.4.1. Terminology 97 3.5.4.2. Pattern of spermatogenic multiplication 100 3.5.4.3. Temporal pattern of young adult spermatogenesis 104 3.5.5. Comparative spermatogenesis 110 3.5.6. General discussion 125 3.6. Oogenesis 127 3.6.1. Introduction 127 3.6.2. Identification of ovarian cell types 127 3.6.2.1. Structural organization of the young adult ovary 127 3.6.2.2. Germinal cell types 129 3.6.2.2.1. Oogonia 129 3.6.2.2.2. Oocytes 131 3.6.2.2.3. Trophocytes 131 3.6.2.3. Somatic cell types 132 3.6.2.3.1. Cells of the ovariole sheath 133 3.6.2.3.2. Germinal epithelium cells 133 3.6.2.3.3. Follicular epithelium cells 133 3.6.2.3.4. Border cells 134 3.6.3. Ovarian development 134 3.6.3.1. Larval development 135 3.6.3.2. Pupal development 139 3.6.3.3. Adult development 143 3.6.4. General discussion 146 4. SUMMARY 149 5. ACKNOWLEDGEMENTS 151 6. SAMENVATTING 152 7. REFERENCES 154 1. INTRODUCTION As the occurrence of large numbersi sa major characteristico f insectpests , insectreproductio n isa ke ytopi ci nth e biological background ofinsec tcontrol . The complexity of this function isreflecte d in the multitude of studies on insect reproduction in its various aspects. One of these aspects is the development of the reproductive cells in the male and female gonads: gametogenesis. In The Netherlands possibilities have been investigated to use the Sterile Insect Method to control the onionfly, Hylemya antiqua (Meigen). Theus eo f this genetic control method, which has been applied successfully in a series of small scale field trials (THEUNISSEN et al., 1973, 1975; TICHELER et al., 1974), involves irradiation and release of flies sterilized by hard X or gamma rays. Irradiation at the wrong stage of gametogenesis may cause adverse effects on the quantity of available sperm cells, the sexual competitiveness and physiolo­ gicalfitness o f the releasedflies. Us e of the correct dose is imperative to avoid unwanted fertility or unnecessary impairment of physiological performance inth eirradiate d individuals.I n ordert oobtai n anoptima l combination ofdose , agean d developmental stagea tth etim eo firradiation , astud y hasbee n madeo f histopathological irradiation effects on male and female reproductive organs and their constituting cells and cell populations (THEUNISSEN, 1976).Th e basis ofthi sstud y isa n investigation of spermatogenesis and oogenesis of H. antiqua from a histological point of view. The histo- and biochemical and genetical aspects of gametogenesis have been omitted deliberately, due to the vastness of this field. Following some remarks on the gross anatomy of the reproductive organs andth eearl ypost-embryonicgona d development,spermatogenesi san doogenesi s are described separately. Spermatogenesis is treated more comprehensively as compared to oogenesis because the latter, and especially the vitellogenesis, has been studied far more frequently in great detail in other insects. After the identification of thegermina l and somatic celltype sinvolve d in gametogenesis, the development of the gonads during larval, pupal and adult life has been described. Apreliminar y comparative description of spermatogenic cell types and structures relates the present work on H. antiqua to other insectspecies . Meded. Landbouwhogeschool Wageningen 76-3(1976) 2. MATERIAL AND METHODS 2.1. INSECT MATERIAL 2.1.1. Bionomics of theonion fly The onionfly, Hylemya antiqua Meigen,belong st o thefamil y Anthomyiidae, the superfamily Schizophora and the suborder Cyclorrapha of the order Dip- tera. KOPPEN (1972) and STORK (1936) described respectively the larval and pupal characteristics of the family Anthomyiidae. Various aspects of the biolo­ gy,rearin g and control of H. antiqua are referred to in a bibliography of SCOTT (1970).Additiona l information isfoun d in ANONYMOUS (1965-1975). Life cycle Theflies emerg e in the spring from pupae which have passed the winter in a state of diapause.
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