Acta anal. 70: 623-640 (1968) Department of Zoology, University of Udaipur, Udaipur, Rajasthan. India Histophysiology and histochemistry of the interstitial gland tissue in the ovaries of non-pregnant marmosets S a r d u l S. G u r a y a Introduction The human ovary at the end of pregnancy shows a marked hypertrophy and development of interstitial gland cells of thecal origin, which do not store the lipid or sterol granules (G uraya [1966a]). However, in the ovaries of non-pregnant women (G uraya [1967a]) and rhesus monkeys (G uraya [1966b]) the corresponding cells of thecal origin attain relatively little hypertrophy and development. The cells of non-pregnancy store secretory, lipid droplets consisting of cholesterol and or its esters, triglyceride and some phospholipid. The interstitial gland cells of non pregnancy are of very transient nature and rapidly revert to the stromal tissue, thus showing no accumulation in the ovary. The current study using histochemical techniques describes the histophysiology, development and nature of interstitial gland cells in the ovaries of non-pregnant marmosets, and compares and contrasts these w ith those of the corresponding cells in the ovaries of women (G uraya [1966 a, 1967a]) and rhesus monkeys (G uraya [1966 b]). In the previous studies, as well as the present one, the term interstitial gland tissue has been used for the cells which develop from the theca interna of atretic (degenerating) follicles. Material and methods Results The m armoset ovary can be easily divided into three zones. The first zone, bounded by th e so-called germinal epithelium , consists of the ovarian cortex which, in its tu rn , consists of the com pactly organized strom al connective tissue (figs. 1-5). It also contains pri mordial follicles, small preantral follicles and m any cords of epithelial cells. The second zone, lying beneath the first zone (ovarian cortex), shows both normal and degenerating antral follicles of various sizes Downloaded by: Vanderbilt University Library 129.59.95.115 - 10/26/2017 9:59:41 PM Ovaries of 43 non-pregnant marmosets (Oedipomidas ordipus) imported from South America were used in the current investigation. Most of them were sexually mature. The histochemical techniques (G uraya [1966a]) previously reported w’ere also employed in this study. 624 Guraya llistopliysiology and histochemistry of the interstitial Figs. 1-14 are photomicrographs from frozen gelatine sections of ovaries fixed in formaldehyde calcium, postchromatcd in dichromate calcium and colored with Sudan black B. Downloaded by: Vanderbilt University Library 129.59.95.115 - 10/26/2017 9:59:41 PM (figs. 1-5). Lying between the follicles is the newly formed interstitial gland tissue which is relatively undifferentiated (figs. 1-5). The third zone contains large patches of interstitial gland cells which show a considerable cytoplasm ic differentiation (figs. 1-4). In the ovaries used for the current study, the follicles did develop and attain a certain size. Follicular atresia subsequently developed 625 Figs. I. 2. 3 and 4, Sections of ovaries from different marmosets, illustrating the variations in the amount of interstitial gland tissue (1G). The ovarian cortex (first zone) shows primordial follicles. The second zone lying beneath the ovarian cortex is occupied by the normal follicles (NF) and regressing follicles (RF) of various sizes. In between the follicles is the relatively undifferentiated interstitial gland tissue derived from the theca interna celLs of atretic follicles. I11 the ovaries of relatively young marmosets, it does not store lipid or sterol droplets (fig. 1). The third zone situated inner to the second zone is occupied by the large patches of hypertrophied interstitial gland cells (1G). The lipids seen in the interstitial gland tissue of the second zone arc absent in the cells of the third zone. The large patches of interstitial gland cells are separated by the stromal tissue. X 20. Acta anat., Vol. TO, Nr. I (1968) 49 Downloaded by: Vanderbilt University Library 129.59.95.115 - 10/26/2017 9:59:41 PM gland tissue in the ovaries of 11011-pregnant marmosets 626 Guraya Hislophvsiology and histochemistry of tlic interstitial and thus there was a failure of follicular growth beyond the critical stage necessary for ovulation. The inhibition of preovulatory growth of the follicle m ay be due to the stresses and strains of captivity. Follicular developm ent beyond a certain stage did not show much im provem ent in the ovaries of those m armosets which were well-kept in the laboratory for several m onths. Both prcantral and antral fol licles can be easily distinguished as normal or atretic follicles by the am ount and nature of lipids present in their granulosa (figs. 1-3), which shows the same histochemical changes as those described for other prim ates (see Introduction). The lipid bodies of healthy fol licles are of heterogeneous appearance. They are sparsely distributed in the oocyte and granulosa. They consist of phospholipid only. The theca interna cells of healthy follicles also show some phospholipid granules. W ith the start of atresia, the lipid bodies consisting of cholesterol and/or its esters, triglyceride and some phospholipid accu- Downloaded by: Vanderbilt University Library 129.59.95.115 - 10/26/2017 9:59:41 PM Fig. 5. Portion of ovary illustrating the distribution of the relatively undifferenti ated interstitial gland tissue between the follicles (healthy and atretic) of varying sizes in the second zone. The interstitial gland tissue is filled with sudanopbilic lipid droplets. The first zone (ovarian cortex) shows primordial follicles and small prcantral follicles. X 36. gland tissue in the ovaries of non-pregnant marmosets 627 Fig. 6. Portion of vesicular follicle, showing the beginning of atresia as illustrated by the accumulation of lipid droplets in the theca interna. The lipid bodies (L2) of regressing granulosa (RG) stain violet in Nile blue. X880. Fig.7. Portion of vesicular follicle illustrating the late stage of degeneration when the granulosa (RG) is greatly crumbled around the antrum. The lipid droplets of atresia have accumulated in the degenerating granulosa cells (RG). The theca interna is idled with sudanophilic lipid droplets. X440. Downloaded by: Vanderbilt University Library 129.59.95.115 - 10/26/2017 9:59:41 PM initiate in the granulosa and theca interna cells (figs.2, 3, 6 and 7). The latter give rise to interstitial gland tissue which shows a m arked hypertrophy and developm ent in the ovaries of non-pregnant m arm o sets. During an increase in the interstitial gland cells, follicles had been developing and were beginning to undergo atresia ((igs.2, 3, 4 and 5). The interstitial gland tissue shows very conspicuous histophysiological and histochemical changes in the ovaries of the same individual. The theca interna cells of a healthy antral follicle show some sparse ly distributed lipid granules which consist of phospholipid. W ith the start of atresia, sudanophilic, coarse lipid bodies begin to accum ulate abundantly in the theca interna cells (figs.6 and 7). These lipids stain pink with Nile blue and red with red Sudan dyes. This indicates that the stored lipids contain triglyceride. They stain intensely for chol esterol and/or its esters. The lipid droplets stain m oderately in acid haematoin followed by a negative reaction in the m aterial fixed in 628 Gubaya llistophysiology and liistocliemistry of the interstitial Fig. 9. Portion of degenerating antral follicle from the ovary of relatively young marmoset, showing the absence of lipid droplets in the cells of thecal origin. The degenerating granulosa cells contain sudanophilic lipids. Compare this figure with figs. 6, 7 and 8. X 440. Downloaded by: Vanderbilt University Library 129.59.95.115 - 10/26/2017 9:59:41 PM Fig.8. Very late stage of follicle degeneration showing the remnants of regressing granulosa (KG) and zona pcllucida (KZP). The cells of thecal origin persist and have nearly filled the space left after the degeneration of granulosa. They are filled with sudanophilic lipid droplets. X 440. gland tissue in the ovaries of non-pregnant marmosets 629 weak Bouin and extracted with hot pyridine. This shows th at they contain some phospholipid, in addition to triglyceride and cholesterol and/or its ester. Such lipids do not accum ulate in the theca interna cells of degenerating follicles in the ovaries of relatively young m ar mosets (figs. 1,9 and 10). Its physiological significance could not bedeterm ined. It is just possible that the theca interna cells of young m arm osets arc not yet adapted to the m etabolism of lipid or steroidal compounds. The granulosa and oocyte finally degenerate and dis appear, leaving behind the theca interna cells which form a patch of irregular shapes around the collapsed follicular cavity (figs.8, 9 and 10). These cells derived from the theca interna cells of degener ating follicles constitute the interstitial gland tissue of the second zone, which is irregularly distributed between the follicles (figs. 1-5). It exhibits poor vascular developm ent. The individual cells are of small size. They do not show much cytoplasmic differentiation. The Downloaded by: Vanderbilt University Library 129.59.95.115 - 10/26/2017 9:59:41 PM Fig. 10. Degenerating antral follicle from the ovary of relatively young marmoset showing the absence of sudanophilic lipids in the cells of thecal origin. The granulosa cells lying inner to the cells of thecal origin show sudanophilic lipid accumulations. X 165. 630 Guraya Histopliysiology and histochemistry of the interstitial I'ig. 11. High power view of the relatively undifferentiated interstitial gland cells of the second zone, which arc tilled with sudanophilic lipid droplets. x440. Downloaded by: Vanderbilt University Library 129.59.95.115 - 10/26/2017 9:59:41 PM cell organelles (m itochondria and Golgi zone) are poorly differenti ated. The most conspicuous inclusions present in their cytoplasm are the coarse lipid droplets (figs.2, 5, 8 and 11), which consist of chol esterol and or its ester, triglyceride and some phospholipid as already described. The third zone of the m arm oset ovary contains large patches of greatly hypertrophied interstitial gland cells, the am ount of which varies in the ovaries of different individuals (figs. 1-4). A transitional area can be found between the second and third zones, owing to very conspicuous changes th at occur in the size and lipids of interstitial gland cells of the second zone (figs.l-A). As the latter approach the third zone, they begin to hypertrophy by exhibiting more cytoplasm ic developm ent (fig. 12). Sim ultaneously, their droplets gradually dis appear from view. After reaching the third zone, they merge com pletely into the previously existing patches of fully differentiated interstitial gland cells (fig. 13) which are well-vascularized, as judged from the presence of num erous capillary loops between them , which are gradually reduced in their developm ent as we go from the third zone toward the second and first zones of the ovary. gland tissue in the ovaries of non-pregnant marmosets 631 The interstitial gland cells of the third zone are polyhedral and show a large vesicular nucleus. But the most striking feature is the lack of lipid accum ulations in their greatly developed cytoplasm (fig. 14). The cell organelles (m itochondria and Golgi zone) are well differentiated. The whole of the cytoplasm including the Golgi zone colors feebly with Sudan black B (fig. 14), indicating the presence of some diffuse lipids. After fixation in Zenker, Carnoy and Bonin, followed by paraffin em bedding, both the cytoplasm and Golgi zone begin to show relatively more coloration with Sudan black, signi fying unm asking of more diffuse lipids in them . After fixation in weak Bonin followed by pyridine extractions, the diffuse lipids dis appear, as judged by a negative reaction with Sudan black. The Downloaded by: Vanderbilt University Library 129.59.95.115 - 10/26/2017 9:59:41 PM Fig. 12. Portion of ovary, illustrating the transitional area between the second (upper) and third (lower) zones. The interstitial gland cells of the second zone are smaller in size: their cytoplasm contains lipid droplets consisting of cholesterol and its ester, triglyceride and some phospholipid. The nuclei (N) arc sudanophobic. As these cells reach the third zone, such lipids disappear; simultaneously the cells increase in size; more cytoplasm is formed. Some sparsely scattered lipid bodies (L.>) are seen among the hypertrophied interstitial gland cells. X880. 632 Gt raya Histophysiology and histochemistry of the interstitial Fig. 13. Showing a large patch of fully hypertrophied interstitial gland cells in the third zone. X 150. Downloaded by: Vanderbilt University Library 129.59.95.115 - 10/26/2017 9:59:41 PM diffuse lipids give negative reactions with various other techniques for lipids. They are due to lipoproteins in the cytoplasm and Golgi zone, which also stain positively with the periodic acid Schiff (PAS) technique. The intensity of the PAS reaction is increased in the m aterial treated with Zenker, Carnoy and Bonin. The PAS-positive reaction continues to appear in the control sections treated with acetvlation. However, the reaction does not appear in the m aterial extracted with pyridine. This dem onstrates that the PAS-positive reaction of the cytoplasm and Golgi zone is again due to their diffuse lipoproteins which are extracted in the hot pyridine. Diffuse lipo proteins are relatively less developed in the cytoplasm of theca in terna cells and interstitial gland cells of the second zone. Both the cytoplasm and Golgi zone give a positive reaction with the m ethylgreen pyronin techniques, which disappears in the control sections treated w ith ribonuclease. This indicates the presence of some KNA in the cytoplasm . The m itochondria are in the form of granules and gland tissue in the ovaries of non-pregnant marmosets 633 Fig. 14. Higher power view ol' fully differentiated interstitial gland cells, showing the nuclei (N). Golgi zone (GZ) and sparsely scattered lipid bodies (L2) of different sizes. The sudanophilic granular mitochondria occur throughout the cytoplasm which colors feebly in Sudan black B. The nuclei ( \ ) are completely sudanophobic. The cells are surrounded by a well marked blood vascularity. X 880. Downloaded by: Vanderbilt University Library 129.59.95.115 - 10/26/2017 9:59:41 PM rods which are sudanophilic, as judged by their coloration with Sudan black. This shows that the m itochondria contain some lipids which react negatively to various other histochemical techniques used for the study of lipids. Sparsely scattered among the hypertrophied interstitial gland cells of the third zone are some lipid bodies (L2) of various sizes (lig. 14); some small lipid granules (L.,) are also seen in the cytoplasm of indi vidual cells. The lipid bodies (L.,) react intensely to Sudan black B and acid hacm atein followed by a negative reaction in m aterial ex tracted w ith hot pyridine. This indicates that the L., bodies consist of phospholipid. The large L., bodies also stain light pink with ¡Nile blue and red with red Sudan dyes, revealing some triglyceride in them , in addition to phospholipid. Of all the fixatives used, the L., bodies are adequately preserved in the m aterial fixed in form aldehyde calcium and postchrom ed in dichrom ate calcium, thus showing the 631 Guraya Histopliysiology and histochemistry of the interstitial Comparisons between the interstitial gland tissue in the ovaries o f women, rhesus monkeys and marmosets The interstitial gland tissue in the ovaries of women (G ukaya [1966a, 1967a|), rhesus m onkeys (G uraya [1966b]) and marm osets is invariably derived from the theca interna cells of degenerating medium-sized follicles and degenerating antral follicles. It shows m any variations in regard to its hypertrophy, developm ent, distribu tion and histochem istry in their ovaries. The cells of thecal origin are relatively undifferentiated in the ovaries of non-pregnant women (G uraya 11967 a|) and rhesus m onkeys (G uraya [ 1966 b |). They store lipid or sterol droplets consisting of cholesterol and or its ester, triglyceride and some phospholipid. The cells of non-pregnancy are of very transient nature, as they rapidly revert to the relatively em bryonic strom al tissue; sim ultaneously they lose their cytoplasm and lipid droplets. This short existence of interstitial gland cells in the wall of degenerating follicles led most of the earlier workers to deny their presence in the ovaries of non-pregnant women and rhesus monkeys. However, the corresponding cells of thecal origin show a m arked hypertrophy and developm ent in the ovaries of non-pregnant m arm osets. The cells are distributed in the second and third zones of the ovary; no such zones develop in the ovaries of non-pregnant women and rhesus monkeys. The cells of the second zone do not show much cytoplasm ic differentiation. In their developm ent and histo chem istry, they resemble the corresponding cells of non-pregnant women and rhesus monkeys. The cells of the second zone in the m arm oset ovary also store coarse lipid droplets in their cytoplasm , which consist of cholesterol and or its ester, triglyceride and some phospholipid. The cells of the second zone, instead of reverting to the strom al elements as described in the ovaries of non-pregnant women (G uraya [ 1967 a|) and rhesus monkeys (G uraya [ 1966b|), gradually transform into large cells which continue accum ulating and leave large patches of interstitial gland cells in the third zone of the ovary. Sim ultaneously, the depletion of lipid or sterol droplets occurs, which is observed in the cells of the second zone. Correspond ing to the hypertrophy of cells and depletion of lipids, there is an increase in the vascularity of the interstitial gland tissue. This sug gests that the relatively undifferentiated cells of the second zone are Downloaded by: Vanderbilt University Library 129.59.95.115 - 10/26/2017 9:59:41 PM necessity of both calcium ions and postchroination for tlicir proper fixation. gland tissue in t he ovaries of non-pregnant marmosets 635 Discussion The interstitial gland tissue of thecal origin shows very conspicuous variations in its am ount, distribution and histochem istry in the ovaries of women (G uraya [1966a, 1967a]), rhesus monkeys (G u ray a [ 1966 b|) and m arm osets. Now we may ask what induces these variations. N elson and G reene [1958] have attributed the m arked hypertrophy and developm ent of thecal cells (interstitial gland cells of the present author) to the influence of placental hormones in pregnant women. According to them , the cells are “ aw akened” by the prolonged and intensive influence of hormones during late ges tation. Now it is reasonable to state that the hormones (gonado Downloaded by: Vanderbilt University Library 129.59.95.115 - 10/26/2017 9:59:41 PM awakened or stim ulated by some factors carried by the well-marked vascularity of the third zone. The accum ulations of interstitial gland cells seen in the ovaries of non-pregnant marmosets do not occur in the ovaries of non-pregnant women and rhesus monkeys. However, their accum ulation begins to occur in the hum an ovary during late gestation (G iraya [1966a], Mossman et al. [1964]). The hyper trophied interstitial gland cells of the human ovaries studied at the end of pregnancy (G i raya [1966a]) and of the third zone in the m arm oset ovary are very similar both morphologically and histochemically, except th at the cells of the marm oset ovary do not show the cytoplasm ic vacuoles described in the relatively m ature cells of hum an ovary (G uraya [1966a]). The cells show much more cyto plasm containing no lipid or sterol droplets. The cell organelles such as the m itochondria and Golgi zone are very prom inent. The cyto plasm including the Golgi zone shows diffuse lipoproteins. There is a meagre developm ent of cytoplasm ic RNA. Some heterogeneous lipid bodies (L.,) consisting of phospholipids are sparsely distributed am ong the hypertrophied cells of the hum an ovary, which are well vascularized. Roth the cytological and cytochemical features suggest that the interstitial gland cells of late gestation in women and of the third zone in the m arm oset ovary are metabolically more active than those of non-pregnant women and rhesus monkeys and of the second zone in m arm osets. In other words, the cells of non-pregnant women and rhesus m onkeys and of the second zone in m arm osets are rela tively quiescent or dorm ant as suggested by 1. the poor developm ent of the cytoplasm and its organelles (m itochondria and Golgi zone), and 2. the accum ulation of lipid or sterol droplets in their cytoplasm . Guraya Hislopbysiology and histochemistry of the interstitial tropins) also stim ulate the corresponding cells in the ovaries of m arm osets, which attain a considerable hypertrophy anil develop m ent; the source of these gonadotropins must be the pituitary, rather than the placenta as the m arm osets used were not pregnant. The presence of some gonadotropins is further supported by the fact that the relatively undifferentiated interstitial gland cells of the second zone hypertrophy only after coming in contact with the well-marked vascularity of the third zone. This indicates th at the cells are stim u lated after they are exposed to a proper am ount of gonadotropins carried by the capillary plexuses of the third zone. The cells of com parable origin in the ovaries of non-pregnant women and rhesus monkeys arc not exposed to such a strong gonadotropic stim ulation and thus rapidly revert to the embryonic strom al tissue. It has been experim entally dem onstrated in the rabbit, rat. and ham ster (G uraya and G reenw ald |19fi4a. b, 19651, G uraya 11967 bj), th at the high levels of both the exogenous and endogenous gonadotropins cause the depletion of lipid or sterol droplets from the interstitial gland cells of their ovaries. After the gonadotropic stim ula tion is over, the cells again store the lipids. These studies have also shown th at the vascularity is correspondingly increased am ong the interstitial gland cells during the strong gonadotropic stim ulation. These experim ents, of course, carried out on different anim als, sup port the suggestion that the lack of proper gonadotropic stim ulation is responsible for the storage of lipid or sterol droplets in the intersti tial gland cells of non-pregnant women (G uraya 11967 a|) and rhesus m onkeys (G uraya (19661>|) and of the second zone in the m arm oset ovary'. The lipid or sterol droplets are not seen in the interstitial gland cells of late gestation in women (G uraya 11966a|) and of the thiid zone in non-pregnant m arm osets, showing their mobilization due to a strong gonadotropic stim ulation to which they are exposed. It will be very interesting to exam ine the lipid changes in the ovaries of women and rhesus m onkeys during the im m ediate preovulatory' period when there is a sudden rise in the levels of luteinizing hormone or interstitial cell-stim ulating hormone. It is generally believed th at when cholesterol is abundantly present in the interstitial gland cells of the ovary', storage is taking place and when the am ount decreases, release of hormone is occurring (see G uraya 11966 a|). According to this view, the interstitial gland cells of non-pregnant women (G uraya [1967a]) and rhesus monkeys (G uraya [1966b]) are acting as storage cells which are relatively' Downloaded by: Vanderbilt University Library 129.59.95.115 - 10/26/2017 9:59:41 PM 636 637 inactive or dorm ant, as already stated. From the various circum stantial evidence it can be said that the relatively dorm ant cells of their ovaries are capable of behaving like active gland cells, provided the proper gonadot ropic stim ulation becomes available. This is clear ly illustrated by the cytological and cytochcmical changes which occur in the com parable interstitial gland cells of late gestation in women and of the third zone in the m arm oset ovary owing to some strong gonadotropic stim ulation. The fully differentiated interstitial gland cells of late gestation in women and of the third zone in the marmoset ovary resemble very closely the interstitial or Leydig cells of the opossum testis in their morphology and histochem istry (G uray A | unpublished observation]). The Leydig cells also show a well-developed cytoplasm containing no lipid droplets. The cell organelles (m itochondria and Golgi zone) are ver\ prom inent. The cytoplasm contains diffuse lipoproteins which have been interpreted as being derived from the extensively develop ed agranular endoplasm ic reticulum described by electron microscopists in the cytoplasm of Leydig cells (C hristensen and F awcett 11961. 1966], C hristensen 119651). The 1GNA is poorly developed in the cytoplasm of Levdig cells. Some heterogeneous lipid bodies con sisting of phospholipid are sparsely distributed am ong the Leydig cells of the opossum testis. The diffuse lipoproteins studied in the cyto plasm of fully differentiated interstitial gland cells of the prim ate ovary, may also be attributed to the developm ent of agranular reti culum, which has been described in the interstitial gland cells of the rat ovary (M uta 11958]). It will be very profitable to make com para tive studies on the submicroscopic organization of interstitial gland cells in the ovaries of prim ates. The presence of abundant granular endoplasmic reticulum in their cytoplasm can be easily ruled out by the fact th at their cytoplasm is poor in RNA, as th a t of Leydig cells (G lraya [unpublished observation]). From the cytological andeytochemical features it is presumed th at the cytoplasm of fully differen tiated interstitial gland cells of prim ate ovary m ust be rich in agra nular endoplasmic reticulum . The biochemical and histochemical studies correlated with morphological findings have clearly shown th at the mem branes of the agranular endoplasmic reticulum in the m am m alian Leydig cells are the site of some enzymes involved m steioidogenesis (C hristensen [1965], Christensen and F awcett [1966]). From the close sim ilarities of the cytological and cytochcm ical features of the interstitial gland cells of both ovary and testis it Downloaded by: Vanderbilt University Library 129.59.95.115 - 10/26/2017 9:59:41 PM gland tissue in the ovaries of non-pregnant marmosets 638 Guraya Histophysiology and histochemistry of the interstitial can he said th at the hypertrophied cells of the prim ate ovary are also steroid-producers. The gland cells of the ovary undergo some rh y th mic changes in their cytology and cytochem istry, which seem to be brought about by the gonadotropins, as already stated. It is well known in m am mals th at the interstitial cells are the main source of testicular androgens (Christensen [1965], C hristensen and F awcett |1966|). The interstitial gland cells of the m am m alian ovary have been considered to form a variety of steroids (estrogens, progesterone and androgens) for which there is no convincing physio logical evidence. However, H i l l i a r d et al. [1963 ] have recently con cluded that the interstitial tissue in the preovulatory ovary of the rabbit is the principal site of progestin synthesis. Acknoivledgements The author wishes to thank all those persons who made the completion of this study possible. The histophysiology and histochemistry of the interstitial gland tissue in the ovaries of non-pregnant marmosets (Oedipomidas oedipus) have been studied with histoehemical techniques. The marmoset ovary can L divided into three zones. Primordial follicles lie in the first zone consisting of the tunica albuginea. Interstitial gland cells occupy the second and third zones. The interstitial gland cells of the second zone are relatively undifferentiated. They are derived from the theca interna cells of degenerating medium-sized and degenerating antral follicles. Their cytoplasm is filled with coarse lipids which consist of cholesterol and or its ester, triglyceride and phospholipid. The third zone (medullary part of the ovary) contains large patches of hypertrophied interstitial gland cells which are gradually derived from the relatively small cells of the second zone. During this transformation, the cells show a consider able cytoplasmic differentiation: the coarse lipid bodies present in llic cells of the second zone disappear from view: the cell organelles (mitochondria and Golgi zone) become very prominent: and diffuse lipoproteins develop throughout the cytoplasm. The cytoplasm of the interstitial gland cells shows some RNA. in addition to lipo proteins. Some lipid bodies (L,), of irregular shape, consisting of phospholipid and little triglyceride, arc sparsely distributed among the cells of the third zone, which seem to be very active physiologically. Interstitial gland cells of the marmoset ovary have been compared and con trasted with those of the rhesus monkey and man. The comparisons suggest that the interstitial gland cells apparently depend upon the gonadotropins for their development, histochemistry and functional integrity. Cytological and cytochcmical similarities found between the fully differentiated interstitial gland cells of the primate ovary and the Lcydig cells of mammalian testis also suggest that the former are steroid sccrctors as the latter, which are now well-known secretors of androgens. Downloaded by: Vanderbilt University Library 129.59.95.115 - 10/26/2017 9:59:41 PM Sum m ary gland tissue in the ovaries of non-pregnant marmosets 639 Résumé Etude hi9tophysiologiquc et liislochimique de la glande interstitielle de l’ovaire chez des Ouistitis (Oedipomidas oedipus) non-gravides. L’ovaire de cet animal pré sente 3 zones. Les follicules primordiaux se trouvent dans la première zone qui forme l’albuginée. Les cellules de la glande interstitielle occupent la 2e et la 3e zones. Celles de la 2e zone sont relativement peu différenciées. Elles proviennent des cellules de la thèque interne de follicules de taille moyenne et de follicules antraux, en voie de dégénérescence. Leur cytoplasme renferme des lipides formés par de la cholestérine et ou ses éthers, des triglycérides et des phospholipides. La 31‘ zone (médullaire de l’ovaire) renferme de grandes plages de cellules interstitielles hypertrophiées, prove nant des cellules relativement plus petites de la 2e zone qui se transforment graduelle ment. Au cours de cette transformation, ces cellules permettent de reconnaître une différentiation considérable de leur cytoplasme: les corps lipidiques rencontrés dans les éléments de la 21' zone ne sont plus visibles: les organclles cellulaires (chondriome, appareil de Golgi) deviennent très nettes, et des lipoprotéines se forment dans toute l’étendue du cytoplasme. Celui-ci renferme en outre un peu d’ARN en plus des lipo protéines. Quelques corps lipidiques (L2) de forme irrégulière, constitués par des phospholipides et des triglycérides en faible quantité, sont répartis parcimonieuse ment dans les cellules de la 3e zone, dont l’activité physiologique parait être considé rable. Les cellules de la glande interstitielle de l’ovaire du Ouistiti sont comparées à celles du singe rhésus et de la femme. Cette comparaison permet de supposer que le déve loppement. l’histochimie et les fonctions de la glande interstitielle dépendent appa remment de l’action de l’hormone gonadotrope. Les ressemblances cytologiques et cytochimiqucs entre les cellules de la glande interstitielle entièrement différenciée de l’ovaire chez les primates, et les cellules interstitielles de Leydig du testicule chez les Mammifères font supposer également que les premières sont des «sécréteurs de stéroïdes» tout comme les secondes, dont la sécrétion d’androgènes est actuellement bien connue. Die Histophysiologic und Ilistochemic des interstitiellen Drüsengewebes in den Ovarien nicht-gravider Seidenäffchen (Oedipomidas oedipus) wurde mit histochemisclicn Methoden untersucht. Das Ovar des Seidenäffcheng kann in 3 Zonen unterteilt werden. Die Primärfollikel liegen in der ersten Zone, die aus der Tunica albuginca besteht. Die Zellen der interstitiellen Drüse finden sich in der zweiten und dritten Zone. Die Zellen der interstitiellen Drüse der zweiten Zone sind relativ undifferenziert. Sie stammen von den Thcca interna-Zellen mittelgroßer degenerierender Follikel sowie degenerierender Bläschenfollikel ab. Ihr Cytoplasma ist angefüllt mit Rohlipidcn, welche aus Cholesterol und oder seinen Estern, Triglyceriden und Phospho lipiden bestehen. Die dritte Zone (Markabschnitt des Ovars) enthält große Flecke von hypertrophierten Zellen der interstitiellen Drüse, die sich nach und nach aus den relativ kleinen Zellen der zweiten Zone entwickeln. Während dieser Umwandlung zeigen die Zellen beträchtliche zytoplasmalischc Veränderungen. Die Rohlipidkör per, die in der zweiten Zone vorhanden waren, verschwinden. Die Zellorgancllcn (Mitochondricn und Golgiapparat) werden sehr auffällig, und diffuse Lipoproteine entwickeln sich im ganzen Cytoplasma. Das Cytoplasma der interstitiellen Drüsen Downloaded by: Vanderbilt University Library 129.59.95.115 - 10/26/2017 9:59:41 PM Zu sum menfassung 640 GiHAYA zellen weist neben Lipoproteiclcn etwas RNA auf. Einige Lipidkörper (L.,) von un regelmäßiger Gestalt, bestellend aus Phospholipiden und etwas Triglyceriden, sind spärlich in den Zellen der dritten Zone verteilt, die physiologisch sehr aktiv zu sein scheinen. Die interstitiellen Drüsenzellen des Seidenäffchen-Ovars wurden mit jenen der Rhesusaffen und des Menschen verglichen und ihnen gegcnübergestellt. Die Vergleiche legen nahe, daß die interstitiellen Drüscnzcllcn hinsichtlich ihrer Entwicklung, ihrer hislochcinischcn Struktur und funktionellen Integrität anscheinend von den Gona dotropinen abhängig sind. Zylologische und zylochemischc Ähnlichkeiten zwischen den vollständig ausdifferenzierten interstitiellen Drüsenzellcn des Primatenovarsund den Lcydigschen Zellen des Säugerhodens legen ebenfalls die Vermutung nahe, daß die ersteren wie die letzteren, von denen man bekanntlich weiß, daß sie Androgene bilden, Steroide sezernieren. References Author's address: Sardul S. Gurtya. Ph.D., Department of Zoology, University of Udaipur, Udaipur, Rajasthan (India) Downloaded by: Vanderbilt University Library 129.59.95.115 - 10/26/2017 9:59:41 PM Christensen , A. K .: The fine structure of testicular interstitial cells in guinea pigs. J. Cell Biol. 26: 911-935 (1965). Christensen , A.K. and F awcett. D.W.: The normal fine structure of opossum testicular interstitial cells. J. Biophys. Biochcm. Cvtol. 9: 633-670 (1961). —The fine structure of testicular interstitial cells in mice. Amer. J. Anat. 118: 551-572 (1966). Gt raya, S.S.: Histochemical analysis of the interstitial gland tissue in the human ovary at the end of pregnancy. Amer. J.Obstet.Gynec. 96: 907-912 (1966a). — A histochemical study of the rhesus monkey ovary. Acta morph.neerl.scand. 6: 395—406 (1966b). Histochemical study of the interstitial gland tissue in the ovaries of non-pregnant women. Amer.J.Obstet.Gynec. 98: 99-106 (1967a). Cytochemical observations concerning the formation, release, and transport of lipid secretory products in the interstitial (thecal) cells of the rabbit ovary. Z. Zellforsch. 83: 187-193 (1967 b). Gt raya, S.S. and Greenwald , G.S.: A comparative histochemical study of inter stitial tissue and follicular atresia in the mammalian ovary. Anal. Rec. 149: 411—434 (1964a). —Histochemical studies on the interstitial gland in the rabbit ovary. Amer.J.Anat. 114: 495-519 (1964b). - A histochemical study of the hamster ovary. Amer. J. Anat. 116: 257-268 (1965). H illiard , .1.; A rchibald , D. and Sawyer , G.: Gonadotrophic activity of pre ovulatory synthesis and release of progestin in the rabbit. Endocrinology 72: 59-66 (1963). Mossman , H.W.; K o erin g , M.J. and F erry . D. jr.: Cyclic changes of interstitial gland tissue of the human ovary. Amer. J. Anat. 115: 235-256 (1964). Ml'TA, T .: The fine structure of the interstitial cell in the mouse ovary studied with electron microscope. Kurume med. J. a: 167-185 (1958). N elson . WAV. and G r e e n e . R .R.: Some observations on the histology of the hu man ovary during pregnancy. Amer. J. Obstet. Gynec. 76: 66-90 (1958). Received November 27th, 1966
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