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The development of the vagina of the domestic fowl Palmer, Violet Elvira 1932

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THE DMELOPMEHT OF THE VAGINA OF THE DOMESTIC FOWL by Violet E lv i ra Palmer A Thesis submitted f o r the Degree of - EASTER OF ARTS i n the Department of ZOOLOGY TEE UBIViSRSIIT OF BRITISH COIOTBIA APPJL, 1958 Aclmo'glerlgment The w r i t e r i s grateful to Dr. C, McLean Praser, Head of the Department of Zoology j, whose encouragement and kindly c r i t i c i s m have been invaluable. She i s also Indebted to Dr. V.S. Asmundson of the Department of Poultry Husbandry, f o r h i s u n f a i l i n g i n t e r e s t and active cooperation. He has given isost generously of h is time and counsel. THE DEVELOPMENT OF THE VAGINA OF THE DOMESTIC FOT/L ISTBODUGTIOH The M&Llerian ducts of the chick have been described (7, 10, 12) as aris ing during the fourth day of embryonic l i f e from a thickened longitudinal band i n the peritoneum ly ing along the outer surface of the mesonephros near i t s attachment to the body wal l , ex-ternal to the Wolffian duct and extending posteriorly to the cloaca* An invagination of the thickened, ep i the l ia l band or tubal ridge takes place at the anterior end of the mesonephros and a groove i s formed. On the f i f t h day the l i p a of the groove fuse to- form a tube which sep-arates from the ridge and which remains open into the body cavity at i t s anterior end to form the coeloiaic aperture of the oviduct. The very short tube thus formed by invagination of the band of epithelium i s situated between the Wolffian duct and. tubal ridge, and ends b l ind-l y posteriorly.. The greater part of the Mftllerian duet i s formed by the backward extension of this tube. I t grows slowly backwards as a so l id rod which gradually becomes tubular and reaches the cloaca dur-ing the seventh day, but does not acquire an opening into the cloaca during embryonic l i f e . The r ight f lnl lerian duct i n the female ceases to develop and degenerative changes take placej but the l e f t duct continues to enlarge and develops as the functional oviduct. HOT?ever, i t has been stated by Gasser, and also by other embryologists who quote M s vork, (4, 7, 9, 10, 12) that t h i s remaining oviduct does not acquire an open-ing into the cloaca u n t i l a f t e r the hen i s s i x months o l d . But- since some I h i t e Leghorns, at the age of 135-150 days, are capable of produc-ing eggs, the present study v,ras undertaken,at the suggestion of Doctor Asmundson, vrith a view to ascertaining the r e l a t i v e age a t the time of establishment of the vaginal o r i f i c e , as i t evidently takes place be-fore the s i x months In early maturing b i r d s , at least.. ' RBvLBg OF THE LIT5RATURE Before proceeding to a description o f . s e r i a l sections made of the oviduct of the chick i n the vaginal region f o r sicroscopieal exas&natlon, I t may be of in t e r e s t to mention what I s stated i n the l i t e r a t u r e about the time of establishment of the vaginal o r i f i c e i n dif f e r e n t oxximals. Kerr (9) i n reference -bo the lower vertebrates i n a general statement sayss "As regards the further development of the Mullerian duct, i t should be noted that i t s completion and opening into the cloaca i s cocmonly delayed t i l l a comparatively l a t e stage - often t i l l a period but shortly before sexual maturity«" In most mammals the vaginal o r i f i c e i s formed before b i r t h (8). In Marsupials each Mullerian duct acquires an opening i n common TTith the Wolffian duct into the-urogenital sinus during embryonic l i f e , as f o r exanple, i n a 36 mm. Dasyurus viverrinus (15). . In adult Marsupials, however, besides tho two l a te ra l vaginae there i s an outgrowth from each Mullerian duct fonaing a median vagina which ends b l i n d l y a short distance anterior to the urogenital sinus* According to H i l l ( 5 ) , the intervening tissue i s ruptured "by the embryo during p a r t u r i t i o n and the opening of the median vagina into the sinus i s thus established. On the other hand, Fletcher (5) considers the opening to be formed by an i n v o l u t i o n of the sinus growing back to meet the cavity of the median vagina when the cavity i s at the point of i t s maximum backward extension. He also found, on examination of sixteen Kangaroos, that the more nearly f u l l grown the animal, the closer the approximation of the b l i n d end of the median vagina to the urogenital sinus. In certain rodents, the vaginal lumen does not extend to the exterior at b i r t h , and the vaginal o r i f i c e i s not established u n t i l sexual maturity.* One hundred female albino mice were examined by Bade and fiosasco (2) who state that!. "The range of the time of the opening of the vagina i s from the twenty-eighth to the f o r t y - n i n t h day of l i f e , Y/ith the median at the t h i r t y - f i f t h day." These workers also made vaginal smears from the mice and found from the appearance of c o r n i -f i e d c e l l s i n the smear, that ovulation usually occurred within twenty-four hours a f t e r the vagina opened. In. r a t s , also, the vagina remains closed u n t i l sexual mat-u r i t y . Long and Evans (11) found that, with r a t s , i n the majority of cases i f the f i r s t ovulation d i d not occur simultaneously with, the establishment -of the vaginal o r i f i c e , i t nevertheless occurred within f i v e days a f t e r . I n the guinea-pig, as described by K e l l y and Papanicolaou (8), the vaginal lumen establishes i t s f i r s t connection with the ex-t e r i o r vdien the female reaches sexual maturity. Previous to that time, the vulvar l i p s are closed and a s o l i d cord of epithelium ex-tends from the termination of the vagina to the- outer l im i t of the vulva. Hie establislinent of the vaginal o r i f i c e i n the toman occurs before-birth (1, 8). The utero-vaginal canal, which, i s formed by the union of the r i g h t and l e f t K u l l e r i a n ducts, i s described (1) as opening i n t o the urogenital sinus by the t h i r d month of embryonic l i f e . During the fourth month the lumen becomes closed again and-near the external o r i f i c e a semicircular f o l d develops which, represents the hymen. The lumen reappears during the s i x t h month and the external opening of the vagina i s permanently established. MATERIAL AID METHOD Female chicks of d i f f e r e n t breeds varying i n age from 45 to 106 days formed the c a t e r i a l f o r t h i s Investigation. With the excep-t i o n of a fevr S.C.?/. Leghorns which were procured from a private breeder, the i n d i v i d u a l l y pedigreed birds were obtained from the Poultry Farm of the University of B r i t i s h Columbia. A portion of the cloacal v;all containing the oviduct was dissected out and placed i n Zenker 1s f l u i d and dehydrated and em-bedded according to the usual technique. S e r i a l cross sections neve made and stained r.ith haematoxylin and eosin. Unfortunately, a large amount of the material obtained at f i r s t proved useless, since i n trying to obtain a portion of the cloaca of suitable size f o r convenient sectioning, i n s u f f i c i e n t tissue v;as dissected out and portions of the posterior part of the vagina were found to have been cut o f f . This was due to the f a c t that when the oviduct reaches the anterior part of the v a i l of the cloaca i t I s Impossible to ascertain I t s course thereafter, and thus, i n order to make cer t a i n of obtaining the entire vagina I t was; found necessary to take extremely large pieces of tissue which made I n f i l -t r a t i o n and sectioning somewhat d i f f i c u l t . DESCRIPTION OF SERIAL CROSS SECTIOHS OF THE VAGIM S.C.Tf. Leghorn 7f4l9g% age 45 daysj weight 298 gnu The upper portion of the vagina Is comparatively w e l l devel-oped vdth high folds proj ecting into a f a i r l y large lumen and with, deep, narrow depressions between. The vaginal lumen, which i s l i n e d with a single l a y e r of columnar epithelium rdth long oval or round nucle i and r e s t i n g on a well-marked basement membrane, i s present f o r only a very short distance, however. Proceeding eaudally, the lumen becomes smaller and smaller u n t i l i t i s reduced to several large s l i t - l i k e openings l i n e d with a single layer of cuboidal or low columnar epithelium, i n the c e l l s of which mitotic figures are numerous. C e l l s i n mitosis are also.abund-ant i n the connective 'tissue stroma surrounding the s l i t s . Gradually the s l i t s become smaller and fewer, but have s t i l l a d e f i n i t e single layer of epithelium* I n places i n the stroma adjac-ent to the s l i t s a dense massing of nuclei i s also apparent. I n these masses the n u c l e i have a somewhat concentric arrangement and surround small areas of pale-staining nuclei containing very few or no granules. This gives the appearance of a hollowing out i n the centre. M i t o t i c figures are abundant i n the surrounding layers of n u c l e i . F i n a l l y the vaginal lumen i s represented by one or two very small s l i t s situated, i n a plate of stroma containing numerous nuclei, and many, c e l l s i n mi-t o s i s . I n some cases, the epithelium l i n i n g the s l i t s i s s t i l l single layered, i n other cases the nuclei are I r r e g u l a r l y massed at the margin of the s l i t , giving the edge a ragged appearance. I n f a c t , i n the l a t t e r instance, the n u c l e i do not surround a r e e l opening, but an area of i n d e f i n i t e , non-staining tissue. C e l l s i n mitosis are present a t the margin. A short distance further p o s t e r i o r l y the s l i t s become o b l i t -erated and the oviduct ends b l i n d l y . L i g ht Sussex K.H.j age 70 days) weight 490 gnu The anterior portion of the vagina i s f a i r l y w e l l developed and the lumen i s long and narrow.. I f i s l i n e d with low columnar, c i l -i a t e d epithelium. The outfoldings are regular, broad, and low, with narrow, shallow depressions between. The vagina i s situated along with the Wolffian duct i n an area of connective tissue stroma containing numerous n u c l e i . Strands of muscle f i b r e and connective t i s s u e , and small infoldings of the cloacal w a l l , separate t h i s area from the l e f t ureter ( F i g . 1 ) . Proceeding f o r a considerable distance caudally, the lumen gradually decreases i n size,; The folds of the vagina become smaller, more Irregular, and p r a c t i c a l l y disappear, from one side of the duct. There i s c e l l u l a r debris i n the lumen (Fi g . £).. As i n the previous case, the oviduct i s f i n a l l y , reduced, to s l i t - l i k e openings l i n e d with epithelium and containing, c e l l u l a r debris consisting of free n u c l e i and finely-granular material in . the lumen (Fig. 5). A single layer of simple cuboids! epithelium containing large,, round n u c l e i i n the centre of each c e l l , l i n e s the small s l i t s f o r the most pa r t , but here and there the c e l l s , have a columnar app-earance. Two or three mitotic figures occur at the margin. There are also a few m i t o t i c figures i n the. adjacent stroma, which i s r i c h l y sup: l i e d with small c a p i l l a r i e s carrying red blood corpuscles. A few eosinophiles and lymphocytes and a f a i r number of red blood c e l l s are scattered throughout the t i s s u e . . In the case of one small s l i t , the. nuclei are i n a single layer on one side, but on the other they are crowding over the margin, have very few granules and are non-staining. Some of these nuc l e i have a shrunken,crenated appearance. In the succeeding sections the s l i t s disappear and one or two m i t o t i c figures are v i s i b l e i n the s o l i d cord of t i s s u e . .Numerous very small c a p i l l a r i e s carrying large numbers of red blood.cells are also apparent. The Wolffian duct and the ureter are s t i l l evident at the b l i n d end of the oviduct, but, i n the case of t h i s b i r d , were not followed further eaudally. Black Orpington % Rhode Island Red #55; age 70 days| weight 444 gm. The vagina has a f a i r l y large lumen and. numerous narrow but well-marked f o l d s . I t i s l i n e d with columnar c i l i a t e d epithelium re s t i n g on a d i s t i n c t basement membrane. The l e f t ureter i s situated at one end and appears i n cross section. This i s the appearance of a considerable number of sections. Then, at the opposite end of the section from the ureter, near the e p i t h e l i a l layer of the cloaca! w a l l , a portion of the o v i -duct w a l l comes in t o view and then the lumen. The portion of the lumen adjacent to the ureter and t h i s new portion next approach and run to-gether to give an extremely long section of oviduct, stretching from the ureter to the margin of the cloaca. The lumen i s quite wide i n places, and the wall i s much folded at the end near the ureter, but folds are absent from the remaining part. This section of the oviduct has the appearance of a longitudinal section, but the ureter, however, s t i l l appears i n cross section* There Is evidently a s l i g h t v a r i a t i o n i n the p o s i t i o n of the vagina as compared with other b i r d s . Tiie vagina gradually decreases i n size., drawing away from the epithelium l i n i n g the cloaca and away from the ureter. I t soon becomes very small and i s l i n e d with low columnar c i l i a t e d epithelium, contain-ing a few c e l l s i n mitosis. I t f i n a l l y ends b l i n d l y shortly before the l e f t ureter opens i n t o the cloaca. Only a few mitotic figures were observed, but an enormous number of eosinophiles "ere present throughout the length of the vagina, both In the epithelium and adjacent stroma. In some places as many as four eosinophil es at once v-ere scmeezed i n between the ^ nu c l e i of the epithelium. Black Orpington #1575; age 91 days} weight 828 gm* The lumen of the posterior portion o f the vagina i s large and the walls are Ruch folded. The folds are high and are separated by deep, side depressions. The duct passes eaudally, adjacent to the Wolffian duct and the ureter, decreasing In si z e f a i r l y rapidly to • three small s l i t - l i k e openings which f i n a l l y fade out and the vagina ends blindly.. The plate of tissue or stroma i s not as d e f i n i t e and does not contain as many nu c l e i i n the region of the b l i n d end as i n the cases described above. There are no mitotic figures. The ureter i s s t i l l present i n cross section and shows no signs o f approaching the cavity of the cloaca. • •• S.G.W. Leghorn #4j age 99 days; weight 819 gm. The vagina Is l i n e d "with c i l i a t e d epithelium r e s t i n g on a well-developed basement nembrane. The nuclei of the epithelium appear to be i n three or four layers. The lumen i s large and the folds i r r -egular. • Proceeding p o s t e r i o r l y , the vagina loses I t s folds and the lumen decreases i n size u n t i l I t becomes very small. Meanwhile, the l e f t ureter has been approaching the cloaca and now opens into i t . The vaginal lumen then diminishes ra p i d l y u n t i l i t i s represented by two t i n y s l i t s containing c e l l u l a r debris i n the lumen, and i s f i n a l l y o b l i t e r a t e d while the ureter can s t i l l be seen entering the cloaca* One m i t o t i c figure i s v i s i b l e i n the area immediately poster-i o r to the b l i n d end of the duct. Black Orpington- x Barred Rock #£104; age 100 days| weight 830 gm. The lumen of the vagina i s f a i r l y large and the folds are regular, broad and low, with, narrow, shallow depressions between. The Wolffian duct i s adjacent'to the vagina and the l e f t "ureter i s j u s t entering the cloaca ( F i g . 4). k short distance posterior to t h i s the lumen becomes smaller. The folds are very low and broad with very shallow depressions. The adjacent stroma, showing a few mitotic figures, i s r i c h l y supplied with c a p i l l a r i e s and there are a few scattered red. blood c e l l s . Proceeding p o s t e r i o r l y , the lumen rapidly becomes much smaller (F i g . 5) and i s l i n e d with low columnar c i l i a t e d epithelium. Then the vagina i s represented by a small s l i t ( F i g . 8) with n u c l e i i r r e g u l a r l y massed i n two or three layers at the margin. 'Three mitotic figures are v i s i b l e i n the adjacent stroma and also a few scattered red blood c e l l s . The lumen i s f i n a l l y re'need to a very t i n y s l i t ( F i g . 7) with nuclei densely, but i r r e g u l a r l y , massed at the margin and wandering across the s l i t . The s l i t disappears and the vagina ends b l i n d l y 460y* posterior to where the l e f t ureter enters the cloaca ( F i g . 8)« In the tissue immediately posterior to the H i n d end tv.-o mitotic figures am v i s i b l e . Tho posterior part of the vagina i s adjacent throughout to the I n f o l d -ing of the cloaca! v?all i n t o which the ureter opens. The l e f t Wolffian duct vras followed some distance posterior to the b l i n d end of the vagina, and was found to end b l i n d l y just-as the r i g h t ureter opened Into the cloaca. S.C.'vT. Leghorn $5% age 106 daysj weight 800 gnu The vagina i s very long and narrow i n cross section and the f o l d s are' extremely low* The l e f t ureter Is approaching the cavity of the cloaca. During the next im sections the ureter enters the cloaca and the lumen of the vagina becomes much narrower, Bith the margins almost tou-ching« Soon the lumen i s reduced to a f a i r l y long s l i t l i n e d \?ith epithelium. About 520yu, , at l e a s t , a f t e r the l e f t ureter has entered the cloaca, the vagina i s reduced to a t i n y s l i t and f i n a l l y ends b l i n d l y . immediately posterior to the s l i t , n u c l e i appear close together In a narrow s t r i p . In places i n t h i s s t r i p the n u c l e i are more loosely associated leaving spaces between. Ro*mitotic figures were observed. DISCUSSION The backward extension of the vagina occurs much aore rapidly i n young birds (45-70 days old) than In older birds (70-106 days o l d ) , as shorai by th'e enormous number of m i t o t i c figures present In the e p i -thelium and adjacent stroma. Even by 70 days the growth Is it<uch slower as evidenced by the small number of c e l l s i n mitosis, while i n b i r d s of 91 to 106 days of age no mit o t i c figures, or at the most, two or three are v i s i b l e . Another evidence of the comparatively rapid growth between 45 and 70 days of age i s that while i n the 45 day old b i r d the vagina has not developed very f a r posterior to the uterus, yet by 70 days i t has almost reached the point adjacent to where -the l e f t ureter enters the cloaca. Between 70 dsys and 91 days there i s not much further growth. In a S.C.T7. Leghorn, 99 daj-s old, the vagina ends b l i n d l y j u s t as the ureter enters the -cloaca. In a 100- day.^  o l d Black Orping-ton z Barred Rock the vaginal lumen extends 460yx posterior to the open-ing of the l e f t , u r e t e r i n t o the cloaca, while the maximum backward ex-tension of the lumen i s reached -in a; 106 day S*CW. Leghorn where the oviduct ends b l i n d l y about 533/*- posterior to the opening of the ureter. The two S.C.vi. Leghorns, aged 99 days and 106 days, respective-l y , i n which the difference i n weight i s i n s i g n i f i c a n t , nevertheless show a considerable difference, as indicated above, i n the development of the vaginal lumen. Eagle and Eosasco (2) found, i n the case of mice which were weighed at the time of establishment of the vaginal o r i f i c e , that there was no c o r r e l a t i o n between the weight of the mice and the time of establishment of the o r i f i c e . B i f f ercncesjln breed, which determine - whether- a b i r d i s early or l a t e maturing, do not ; seenr to-kavs any e f f e c t on the development of the vaginal lumen during the period observed. The degree of backward extension of the lumen appears to be correlated with the age of the b i r d regardless of i t s breed. There i s a gradual development of the lumen i n extent, during the 45 to 108 day period, through a S.C.W. Leghorn of 45 days, a 70 day Light Sussex and a 70 day Black Orping-ton x Rhode Island Red, each shor/ing a si m i l a r degree of development, a 91 day Black Orpington, a 99 day S.C.W. Leghorn, a 100 day Black Orpington x Barred Rock, and f i n a l l y , a 106 day S.C.?;. Leghorn. As regards the actual process of formation of the vaginal lumen, there i s very l i t t l e data given i n the l i t e r a t u r e . The vagina i s usually spoken of as "acquiring" a lumen. K e l l y and Papanicolaou (8) r e f e r to the development of the vaginal lumen i n the human foetus as a vacuolization process. I t i s also s i m i l a r l y described by other v/riters ( l , 6) as due to the breaking down of the central c e l l s of the duct and the arrangement of the peripheral ones Into epithelium. In b i r d s , the process of development of the lumen appears to resemble that described as taking place I n the human. The vagina narr-ows dorm to a small s l i t - l i k e tube. This narrow portion at the b l i n d end very gradually elongates, growing back into the thickened plate of stroma and enlarges by the growth of the epithelium. At the growing t i p , the n u c l e i become arranged i n concentric masses surrounding a small area of c e l l s which becomes i n d e f i n i t e and non-staining tissue and f i n a l l y breaks down forming a small s l i t . From the nuclear layer at the margin of the s i l t s simple cuboidal or columnar epithelium i s formed. As mentioned above, mitotic figures are numerous at the mar-gin of the s l i t s i n the 45 day old b i r d , shoeing the epithelium to be rapidly growing at t h i s stage. The t i n y s l i t s vrith nuclei densely but i r r e g u l a r l y massed at the margin, and i n some cases wandering across the opening, the some-what ragged appearance of the margin, the c e l l u l a r debris'consisting of scattered pale-staining n u c l e i and i n d e f i n i t e granular material i n the lumen of the posterior end. of the duct, are a l l evidences of the breaking dovm of the tissue enclosed by the layer of n u c l e i . In the human the breaking dovm of the central c e l l s of the vagina i s described (8) as s t a r t i n g i n the d i s t a l p ortion of the vagina and proceeding c r a n i a l l y . In the chick, however, the lumen appears to form almost simultaneously i d t h the development of" the l i n -ing epithelium. This seems to be si m i l a r to. what occurs i n the anter-i o r portion of the human Mullerian duet (8). The numerous c a p i l l a r i e s contained In the thickened cord of tissue i n which the- vagina, develops, r i c h l y supply the rapidly growing tissue with blood. The large number of eosinophiles observed i n the case of one of the b i r d s was probably not connected with the process of formation of the lumen, but due to some l o c a l i n f e c t i o n of the o v i -duct. I t i s evident from the foregoing that the main object of t h i s study has not been attained, namely, the finding of the r e l a t i v e age of the chick a t the time of establishment of the vaginal o r i f i c e . But, due to the time of year i t i s impossible, at present, to obtain older birds f o r further investigation.. However, something has been learned of the nature of the development of the vaginal lumen and i t has been d e f i n i t e l y ascertained that the vagina does not open into the cloaca before the b i r d i s 106 days old . Since, then, as shown by a review of the l i t e r a t u r e , i n those animals i n which the vaginal o r i f i c e i s not established before b i r t h , the vagina does not open u n t i l the attainment of sexual maturity, and since the rate of backward extension of the lumen i s r e l a t i v e l y very slow i n birds 70 - 106 days of age, i t i s quite probable that the vagina does-not develop an opening i n t o the cloaca u n t i l the b i r d also i s sexually mature, that i s , i n Leghorns, at about 1S5 150 days' of age, end correspondingly a l i t t l e l a t e r i n l a t e maturing birds* - -16 — REFERENCES TO. LITERATURE 1. Bailey, F.R., and M i l l e r , A.11. Text-Book of Embryology, 5rd ed., Hew York, 1920, pp. 413-415. 2. Engle, E.T., and Rosasco, J . The Age of the Albino Mouse at Normal Sexual Maturity. Anat. R ec, V o l , 36, 1927, pp. 583-388. S. Fletcher, J . J . On Some Points i n the Anatomy of the Urogenital Organs i n Females o f Certain Species of Kangaroos. — Part I I . Proe* Linnean Soc. lew South Wales,. Vol. 8, 1885-1884, pp. 6-11. 4. Gasser, E. Beitrage sur Entwickelungsgeschichte der A l l a n t o i s , der Mullersehen Gauge und des Afters. Frankfurt a.M., 1874, (quoted from 7, 10, 1?.). 5. H i l l , J.P. Contributions to the Morphology and Development of the Female Urogenital Organs i n the l a r s u p i a l i a . Proc. Linnean Soc. New South Wales, V o l . 24, 1893, pp. 42-80. 6. Keibel, F., and M a l l , F.P. Manual of Human Embryology, V o l . 2, Philadelphia, 1912, pp. 911-952. 7. K e l l i c o t t , W.E. Chordate Development. Hew York, 1915, pp. 560-561. - 17 -8. K e l l y , G.L., and Papanicolaou, G.N. The Mechanism of the P e r i o d -i c a l Opening and C l o s i n g o f the V a g i n a l O r i f i c e i n the Guinea-Pig. Am, Jour. M a t . , Y o l * 40, 1927, pp. 587-401. 9. K e r r , J.G. - Text-Book of Embryology, V o l . 2, London, 1919, p. 245 and p. 554. • • •" * 10. L i l l i e , F.R. -The'Development o f the Chick. 2nd ed., Hew York, • • 1927, pp. 400-405. 11. Long, J.A., and Evans, H. IfcL. The Oestrous Cycle i n the Rat. U n i v e r s i t y o f C a l i f o r n i a Memoirs, V o l . 6, B e r k e l e y , 1922, pp. 50-52. 1-2. McEwen, R.S. A Text-Book .-of-Vertebrate Embryology, Her; York, 1923, p. 438. 13. Van den Eroek, A.J .P. Zur Entv.lckelung der Geschlechtsstrange und Geschlechtsgange b e i den B e u t e l t i e r e n . Anat. Anz., Bd. 28, 1906, pp. 579-594. Explanation of Figures Light Sussex H.N.; age 70 days.' Gross section showing vagina and Wolffian duct situated In an area of stroma and separated-from the l e f t ureter by strands of muscle-fibre and connective tissue and an-infolding of the cloaca! Trail. Haematoxylin and so nisi. X 48. Same. Cross section of oviduct posterior to figure 1, showing vagina narrowing down .as i t approaches the b l i n d end, and closing over as shown by the two s l i t s at the upper r i g h t . Sote the c e l l -u l a r debris i n the lumen. The Wolffian duct I s v i s i b l e at the lower r i g h t . Haesatojylin and eosin. - X 100. P L A T E I . PLATE 2 Explanation of Figures Same. A few sections posterior to figure 2* The vaginal lumen, which i s almost obliterated, i s represented by one large s l i t - l i k e opening and four smaller s l i t s . The Wolffian duct i s at the lower r i g h t . Bote the dense plate of stroma and also the numerous cap-, i l l a r i e s . Haematoxylin and eosin. X. 100. Black Orpington x Barred: Bock #2104? age 100 days. Gross section of cloaca! w a l l showing the vagina with a f a i r l y large lumen and -broad, low f o l d s . At the upper l e f t , the l e f t ureter can be seen entering the cavity of the cloaca. The Wolffian duct i s at the lower ri g h t . Haematoxylin and eosin. X 40. PLAT E H . PLATE 5 Explanation of Figures Same, A few sections posterior to figure 4.- The vaginal lumen has become much smaller,,. The Wolffian duct i s at the lower right* Haematoxylin and eosin. X 80. Same* The vagina i s represented by a small s l i t - l i k e opening. The nuclei are' i r r e g u l a r l y massed i n two or three layers at the margin* Haematoxylin and eosin. X 500* PLATEH. PLATE. 4 Explanation of Figures Same. The vagina i s reduced to a very small s l i t with nu c l e i densely "but i r r e g u l a r l y massed at the margin and wandering across the s l i t . Haematoxylin. .and eosin. X 650. Same. Cross section at the b l i n d end of the vagina 460^ posterior to figure 4. Bote the Wolffian duct at the lo-right. Haematoxylin and eosin.. X 80. 

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