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TEE UNIVERSITY of OKLAHOMA GRADUATE Coiiiege ' A TEE UNIVERSITY OF OKLAHOMA GRADUATE COIIiEGE ' A COMPARATIVE HISTOLOGICAL AND HISTOCHEMICAL STUDY OF THE ADRENAL GLANDS OF NATIVE RABBITS A THESIS SUBMITTED TO THE GRADUATE FACULTY ±n partial fiolflllment of the requirements for the degree of DOCTOR OF PHILOSOPHY BY I. ERNEST GONZALEZ Oklahoma City, Oklahoma 1955 A COMPARATIVE HISTOLOGICAL AND HISTOCHEMICAL STUDY OF THE ADRENAL GLANDS OF NATIVE RABBITS APEROVED BY THESIS COMMIT' ACKN0WEE33GEMENT The writer wishes to express his profound appreciation and sincere thanks to Dr. Kenneth M. Richter, Department of Anatomy, University of Oklahoma Medical School, for his valuable time, coopera- I _ ition, helpful criticisms, and timely suggestions during the course of j I this investigation; to Dr. Ernest Lachman, Chairman of the Department of Anatomy, for his encouragement and cooperation; to Dr. Garman Daron, Professor of Anatomy, for his many helpful suggestions % and to the University of Oklahoma for a University scholarship. Many other persons have cooperated indirectly in making this investigation possible, and the writer would like to acknowledge also the assistance of Dr. C. Lynn Hayward and Dr. D. Eldon Beck, Depart­ ment of Zoology, Brigham Young University, for procuring and identify­ ing many of the native rabbit species; of Mr. Ernest Reiser for his advice during the preparation of the graphic models; and of Mr. Neil Woodward for his assistance with the photomicrographic reproductions. ill TABLE OF CONTENTS Page CHAPTER I 1 Introduction CHAPTER II Materials and MetHods CHAPTER III Observations .............................. 7 Ocbbtona princeps ............»...... ...... 7 Pexicapsular tissue^ capsule, and stroma 7 Vasculature . ......... ............. 8 Innervation............... .... ........ 9 Cortex ........ , . ... ........ ... .. 9 Zona glomerulosa....... ............ 9 Zona fasciculata .............. 12 Zona reticulari s ................... 13 Medulla ...... ........................ l6 Lepus califomicus ..... ................ 19 Perleapsular tissue, capsule, and stroma 19 Vasculature .... .............. 20 Innervation ..................... ...... 21 Cortex .... .................... ........ 22 Zona glomerulosa ...... .......... 22 Zona fasciculata ........... ....... 23 Zona reticularis .......... 2 i+ Medulla ; ............ ............ 25 Lepus tovnsendii ....... .......... 28 Pericapsular tissue, capsule, and stroma 2 8 Vasculature .......... ......... 29 Cortex ....... .......... 30 Zona glomerulosa................... 30 Zona f a s c i c u l a t a...... ..... ...... 32 Zona reticularis .......... 33 Medulla ...... ................. 33 iv riage Lepus americarius 34 Pericapsular tissue, capsule, and stroma ......... 34 Vasculature 35 Innervation .................... 35 Cortex ......... 35 Zona glomerulosa .......... 35 Zona fasciculata. and zona reticularis ............ 36 Medulla --- ...------ 37 Sylvllagus nuttallli ...... 37 Pericapsular tissue, capsule,- and stroma ........ 38 Vasculature ..... 38 Innervation ........ 39 C o r t e x - ...... ... .... 39 Zona glomerulosa ....... 39 Zona fasciculata.................. ....... 42 , Zona reticularis ..... 43 Medulla ...................... ■Ml- IHAPTER IV Discussion . - 46 :h a p t e r V BuTtmtary ...... ..... ............ 81 A. Observations .... 8l B» Belationsliip to other speciès .................... 87 [ME T E K VI Conclusions ...... 91 LITERATUEIE C U E D 9^ PLATES AND LEGENDS ... 10° LIST OF ILLUSTRATIONS Page p l a t e I ................. 100 PLATE II ...... ............................ ..... ......... 101 j PLATE III Abbreviations ......... 102 Explanation, of Plates .......... 102 Explanation, of Figures 1^ 2-, 3> and ^ .. .... .. ... .... 102 PLATE IV Explanation of Figures 5^ 6 , 7> and 8 ............... 103 .PLATE V Explanation of Figures 9f 10, 11, and 12....... .......... 104- PLATE VI Explanation of Figures 13, l4, 15, and l6 ..... 105 PLATE VII Explanation of Figures 17, l8 , 19, and 20 ... 106 ITL A COMPARATIVE HISTOLOGICAL AND HISTOCHEMICAL STUDY OF THE ADRENAL GLANDS OF NATIVE RABBITS I CHAPTER I I i I INTRODUCTION | In recent years, and particularly in the last decade, a great | ideal of research has heen centered on the morphology and especially , the function of the adrenal gland. The vast and voluminous literature i I jin all fields of adrenal research clearly indicates a need for a better i understanding of the processes involved in the biosynthesis and elabor- i at ion of hormones (Dorfman and Unger, ’5^) 7 the physiological role ! I j I (Hartman and Brownell, *4$), and the morphological changes that acconçiany Inormal and pathological adrenal stress (Selye, '46). 1 : ' I In order to evaluate properly and integrate the physiological. jand morphological changes accompanying a variety of stresses, it becomes ! jimperative that we acquire a clearer understanding of the normal paren- jchymal arrangement, cyto-architecture, and histochemistry in different I mammalian species. Differences among species seem to be both physiological and morphological, with some indication that they may extend down to the various strains of any one species or subspecies (Bourne, *49). The variations in the histology, cyto-architecture, and histochemistry in ithe -various mammalian orders is quite apparent, tut few, if any, conrprel- 1 I hensive studies have heen made to determine the degree of variation amohg families, genera, and species of any one mammalian order or group of closely related mammals (Bachman and Schanrer, ’5^) » Recent hooks hy Bourne ( '^9), Hartman and Brownell ('49), and Bachman and Scharrer (’54), in which the literature has heen admirahly reviewed, clearly reveal our lack of knowledge in conparative adrenal histology. Since the mammalian order Lagomoipha contains only two families and relatively few genera, it affords a splendid opportunity for a com- ! : , i parative histological study of the adrenal gland within a group of i i closely related mammaJLs. A review of the literature indicates that the I ! j adrenal histology of Lepus cuul cuius (the domesticated rahhit) is w e H - I known, and this information will form the hasis fof many comparative I remarks. I The purpose of this investigation will he to make a comparative I study of the histology, cyto-architecture, and histochemistry of thé I adrenal gland of the following species within the order Lagomorpha; Ochotona princeps (Rocky Mountain pika), Lepus townsendii (mountain hare), Lepus «mericanus (snowshoe rahhit), Lepus califomicus (desert jack rahhit), Sylvilagus nuttallii (cottontail rahhit), and Sylvilagus idahoensis (pigmy rahhit). Although the nature of this study is for the most part morpho­ logical, it is hoped that hy studying and conparing mammalian species which hitherto have not heen investigated, we might broaden pur general folowlëdge of the adreiml histology 'and mate morë^cômpatlhle p r ê ^ l l ï h ^ physiological and morphological views which mi^t hring us to a closer Understanding of the adrenal function. CHAPTER II MATERIALS AND METHODS The observations made in this study have been based on the | adrenal glands of approximately two hundred specimens of the following | species within the order Lagomorpha: 39 Ochotona princeps (pika), 26 Lepus townsendii (mountain hare), 23 Lepus americanus (snowshoe rabbit) 63 Lepus califbrnicus (jack rabbit), 37 Sylvilagus nuttallii (cotton- j tail rabbit), and 6 Sylvilagus idahoensis (pigmy rabbit). The various ! species were identified by Doctors C. lynn Hayward and D. Elden Beck, j I Department of Zoology, Brigham Young University. ! In order to minimize possible sources of error, specimens for {this study were selected only if they appeared mature and healthy. The I rabbits were collected at monthly intervals over a period of two years, land only those specimens collected during January, February, and "March appeared to be sexually active. Specimens from both sexes were seleoted for this study. All the native rabbits were shot, and those which did not die instantly were immediately killed by a sharp blow on the head. The adrenals were removed with minimum manipulation and immediately fixed by immersion. One gland, in toto, was fixed in Bouin’s fluid. In order to minimize possible artifacts of fixation, the other gland was cut into two equal parts, and one hal f was fixed in Zenker-formal, the other in. the following modificatïônTof Worchester' s'^fixativeT Saturated mercuric chloride in 10^ formalin 10 parts Absolute ethyl alcohol 1 0 parts ! I Glacial acetic acid 1 part After the gland had remained in ¥orchester’s fixative for j twenty-four hours, the tissue was transferred to .alcohol, A number | of glands were also fixed in Severinhaus ' modification of Champy's | fixative I other glands were postchromated in a 3 per cent solution of potassium bichromate. Fresh adrenals or glands fixed in either hufferedj neutral formalin or Baker's calcium-formal were employed for the histo- | ; chemical studies of the lipids. ' | Glands used for architectural reconstructions were embedded in i i ! i paraffin, sectioned at 10 micra, and mounted serially. For critical | fcytological studies, glands were sectioned at 5 micra and the majority I were mounted serially,. Glands used for histochemical studies were ifrozen «--pfl sectioned or were embedded in polyethylene glycol wax after the method of Rinehart and Abu-Haj ( ' $l). Using the latter technique, sections 2 to 5 micra were easily obtained. Most sections were stained routinely in hematoxylin and eosin, ! Mallory^s or Masson's stains,
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