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A Histochemical Identification of the Cellular Sites of Androgen Synthesis in the Hypertrophied Right Gonad of the Leghorn Poulard

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A Histochemical Identification of the Cellular Sites of Androgen Synthesis in the Hypertrophied Right Gonad of the Leghorn Poulard Loyola University Chicago Loyola eCommons Dissertations Theses and Dissertations 1964 A Histochemical Identification of the Cellular Sites of Androgen Synthesis in the Hypertrophied Right Gonad of the Leghorn Poulard James Edward Woods Loyola University Chicago Follow this and additional works at: https://ecommons.luc.edu/luc_diss Part of the Anatomy Commons Recommended Citation Woods, James Edward, "A Histochemical Identification of the Cellular Sites of Androgen Synthesis in the Hypertrophied Right Gonad of the Leghorn Poulard" (1964). Dissertations. 738. https://ecommons.luc.edu/luc_diss/738 This Dissertation is brought to you for free and open access by the Theses and Dissertations at Loyola eCommons. It has been accepted for inclusion in Dissertations by an authorized administrator of Loyola eCommons. For more information, please contact [email protected]. Copyright © 1964 James Edward Woods A HISTOCHEMICAL IDENTIFICATION OF me CELLULAR SITES OF ANDROGEN SYNTltESIS IN THE HYPERTROPHIED RIGUT GONAD OF THE LEGHORN POULAJU) a\\CH SCHO "-." 0 C!) LOYOLA (' UNIVERSITY Of: MED\C\~~ by J..... Edward Wood. A Dissertation SUbaitted to tbe Faculty of tbe Graduate School of Loyola University 1n Partial Fulfillment of the Require.-nts for the Degree of Doctor of Philosophy June 1964 ACKNOWLEDGEMENTS I wish to express ~ sincere appreciation to Dr. Lincoln V. Ooma for suggesting the probl_, indoctrination into the vagaries of sex. biology and for his patience throughout the exteaded period of this investiaation. To Or. Leslie A. emaert for many iDterestlng discussions on the historical aspects of lipid histochemistry and for advice and counsel on a mul. titude of aatters pertaining to the dissertation go my heartfelt thanks. Dr. Jose lannakaril aided in performance of sinistral o.ariectoa!e, on very young biNs. Appreciation i. hereby rendered. Thanks are extended to Dr. Louise Smelte, Histological Technician in the Anatomy ~art... t, for SUIlest10ns OR histological matters. Gratitude for the loan of photop-aphlc eqtd,p1M11t and for export advice on photographic techniques 11 extoded to Dr. Robert C. Thoaes, DePaul University. VITA James Edward Woods was born in Chicago, Illinois, on May 7, 1926, and was graduated frOlat Steimaet& Hilh Sehool in June, 1944. Prom July, 1944, through Sept81lber, 1946, he served in the U. S. Array. In 1951 he received the B. S. Degree in ZooloIY frOIl the University ot Illinois. The following three years he worked at the Vio 81n Corporation, Monticello, Illinois, and the Research Institute of Michael RH,e Hospital, Chicago. In Septellber, 1954, the author began h1s graduate studi•• at DePaul University, receiving the M. S. Degree in Zoology 1n Aulust, 1957. In October, 1957, he enrolled in-the AnatOll), Departaent of the Stritcft School of Medicine of Loyola University. During the period spent In the Anatomy Depar~nt he has helo the following fellowshipst StanclaN Oil "ellow.hip (1951-1960), Loyola University Teaching Pellowship (1961), and Natioaal Institute of Health Predactoral Pellow­ ship (1962-1964). The author is a member of the American Association for the Advance­ ment of Science and the American Society of Zooloalsts. ABSTRACT Although the vertebrate ,oRad is known to be the primary source of the sex hormone., the cells within the.e orlans responsible for thoir production ~ave not been conclusively demonstrated. In particular, the cellular sites of ~droaen synthesi. have not been definitely identified. Most of the experiaental evidence bas implicated the interstitial ~.11s of Leydi. as the principal site. of androgen synthesis in the testis (Albert_ 1961), but there is also evldeace that the se.iniferous tubule•• ay ~onstitute a secondary source of the ..Ie sex hormones. In the ovary the evi­ ~ence indicate. that tbe .-dullaroy interstitial cells are responsible for the production of androgens (Hill, 1962). The purpose of the pre.ent investigation was to identify by histo­ ~hellical Hans the androgen-producing ceUs of the vertebrate gonad, with par­ ~icular reference to tbe testis-like, hypertrophied, rudimentary right gonad of :the sinistrally avarieet_Iled, single-cOllb, light-brown Leghom fowl. This gooad is known to secrete andro,enic hormone. as shown by the effects on be. bayior and on target organs, particularly comb and wattles (Dom., 1921. 1929). However, then is no preche Inf'oraation as to the particular cells within this DTlaD responsible fOT the syntheSiS of the.e hormones. Since the pTe,ently available histochemical techniques for the identification of steroids lack specificity, a new .ethod fOr the identification pf androgens was developed for use in this inv.stigation. It is a .edified fluorescent antibody technique, which is capable of identifying steroids -in !!!!_ The fluorescent anti-steroid sera used in this investigation was prepared against testosterone, and gives a minimum of cross-reaction with other steroids_ Results following -.ployaent of tbis technique have been compared with those obtained from the use of the steroid dehydrogenase method (Watten­ ~erg, 1958). This latter .ethod indicates cellular sites of steroid synthesis Iby vi!lualldn, hist.ologically one of the enzymes (,AS .. 38-hydroxysteroid dehydro- genase) involved in the biogenesis of the steroid hormones. J~ever. it pro­ ~ides only iruUrect evidence tor steroid synthesis, since the presence of a ~ingle enzyme in the biosynthetie pathway does not insure successful synthesis, let alone release of formed hormone (Lobel !l~ 1962). Also, it does not pro­ ~ide inforaation as to the character of the steroid hormone end-product. Histocheaical examination of the testiS-like, hypertrophied, rudi. ~entary right lonacia of brown Leghorn poulards, correlated with the androgen­ eondi tioned chu,e. observed 1n the sexual cbaracteTs of such low I, indicate ~hat the cellular sources of the androgenic steroids produced by this gonad are the interstitial cells as well as the .adultary cords and tubules. The testes and ovaries of nonu.l domestic fowl and albino rats, as ~.ll as the teste. of a dog. were al,o examined histochemically. Positive Toac- tlons were observed in the interstitial cells of Leydig and all cellular ele. Dents of tbe seminiferous tubules in the testis and in the medullary interstl- tial cells of the ovary. TherefoTe, it Is considered that all of these cellular site. are capable of androgen synthesis. TASLE Of CONTENTS Page INTRODUCTION •••••••••••••••••••••••••••••••••••••••••••••••• 1 REVIEW 0' LITERATURE •••••••••••••••••••••••••••••••••••••••• Biology ••••••••••••••••••••••••••••••••••••••••••••••••••• 4 T.stis ••••••••••••••••••••••••••••••••••••••••••••••••• 4 Ovary •••••••• ••••••• .......... •••• •.••••••••••••••••••••• 11 Hypertrophithi. Rudirumta1')' Right Gonad ••••••••••••••••• 13 Bloch.mistry •••••••••••••••••••••••••••••••••••••••••••••• 11 Histochemistry •••••••••••••••••••••••••••••••••••••••••••• 21 MATBRIAIS AND MEllIODS ••••••••••••••••••••••••••••••••••••••• 2S OBSERVATIONS •••••••••••••••••••••••••••••••••••••••••••••••• 38 H1stology of the Hypertrophied Right Gonad •••••••••••••••• 39 Histochemistry of the Hypertrophied Right Gonad ••••••••••• 41 Steroid Dehydrogenase Methed ••••••••••••••••••••••••••• 41 Fluorescent Antibody Method •••••••••••••••••••••••••••• 43 Validation •••••••••••••••••••••••••••••••••••••••••• 43 Observations •••••••••••••••••••••••••••••••••••••••• 44 Hi,tocluaistry of No1'tl81 Gonads ............. •••••••••••••• 46 Gonads of Leghorn Powl ••••••••••••••••••••••••••••••••• 46 Mammalian Gon.o. ••••••••••••••••••••••••••••••••••••••• 47 Pa,o DISCUSSION •••••••••••••••••••••••••••••••••••••••••••••••••• 49 SUMMARY ~~O CONCLUSIONS ••••••••••••••••••••••••••••••••••••• S6 LITERATURE CITED •••••••••••••••••••••••••••••••••••••••••••• 58 TABLES •••••••••••••••••••••••••••••••••••••••••••••••••••••• 11 PLATES •••••••••••••••••••••••••••••••••••••••••••••••••••••• 17 INTRODUCTION The gonads of higher vertebrates are the primary source of sex hormones. however, the cells within these organs responsiblo for the synthesis of these hormones have never been conclusively demonstrated. In particular, the cellular sites of male sex hormone production have not been definitely identified. A considerable accumulation of experimental data has implicated tht interstitial cells of Leydig at the principal sites of androgen synthesis in the testis (Albert, 1961), but there is also evidence that the seminiferous tubules may constitute a secondary source of this hormone (Nalhandov, --et aI, 1946). In the ovary the evidence indicates that the interstitial cells lire responsible for the production of aadNlons (Hill, 1%2). The purpose of the present investigation was to identify by biste- chemical .eans the androgen-producing cells of the vertebrate gonad, with part!- cular reference to the hypertrophied, rudimentary right gonad of the sinistrally ovariectomized, lioglo-cOllb, light-brown Leghorn fowl. This gonad is known to secrete androgenic hormones as shown by the effects on behavior and on target Iorgans, particularly comb and wattles (i)Ollll, 1927, 1929). However, there is no ~recise information as to the particular cells within this organ responsible for ~he synthesis of these hormones. Since the presently available histochemical technic~ues for the identification of steroids lack specificity, a Rew method for the --in situ iden- tifieation of androgens was developed for use 1n this investigation. It is a modified fluorescent antibody technique, and takes advantage of the fact
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