Get access to all handy features included in the IVIS website
- Get unlimited access to books, proceedings and journals.
- Get access to a global catalogue of meetings, on-site and online courses, webinars and educational videos.
- Bookmark your favorite articles in My Library for future reading.
- Save future meetings and courses in My Calendar and My e-Learning.
- Ask authors questions and read what others have to say.
The Ovary
Get access to all handy features included in the IVIS website
- Get unlimited access to books, proceedings and journals.
- Get access to a global catalogue of meetings, on-site and online courses, webinars and educational videos.
- Bookmark your favorite articles in My Library for future reading.
- Save future meetings and courses in My Calendar and My e-Learning.
- Ask authors questions and read what others have to say.
Read
Foreword
The study of the macro and microanatomy of the female genital tract is of great importance to the clinician. Understanding the normal anatomical position and morphology of all parts of the reproductive tract and related structures during all phases of reproduction is the key element for a thorough examination and evaluation of this system by rectal palpation and/or ultrasonography. Also, the knowledge of the histology of these organs - and in particular that of the uterus -enables biopsy techniques to be used as a method for breeding soundness examination. This chapter covers the essentials of reproductive tract anatomy and histology in the female which will aid the reader to better follow the methodology described for the breeding soundness examination.
Morphology and Position
In the dromedary, the ovaries are suspended by the mesovarium at the level of the sixth or between the 6th and 7th lumbar vertebra, about 6 to 7 cm cranial to the crest of the ilium (Figure 5.1).(107, 108) Theovary js located at about 36 cm from the opening of the vulva. The position of the ovary is subject to great variations depending on the physiological stage - especially pregnancy. It becomes more ventral and is pulled forward with advancing stages of pregnancy. The left ovary is usually more cranio-ventral in position than the right ovary.(44, 76, 105, 107)
The ovary is attached to the broad ligament by a well-defined, strong ligament (ovarian ligament or proper ligament) which extends from the hilus of the ovary to the tip of the corresponding uterine hom (Figure 5.2). This ligament originates at mid-distance between the lateral border of the uterine hom and the ovary.(96, 105, 107) Both ovaries are enclosed within a fold of the mesosalpinx known as the ovarian bursa or bursa ovarii (Figure 5.3 and 5.4).(30, 31, 48, 76, 87, 96, 106) The ovary can easily be exteriorized from its bursa. Its general appearance and size varies according to activity and age of the animal. It is small and has a smooth and glistening surface in prepubertal animals (Figure 5.5). Several raised small vesicles can be seen throughout the surface of the ovary and correspond to follicles (2-5 mm in diameter), giving the ovary a granular appearance (Figure 5.6).(31, 76) The ovary increases in size and becomes more irregular and lobulated in older animals (Figure 5.7). This lobulation increases with increased number of previous ovulations or pregnancies and is due mainly to the presence of old corpora albicantia.(76) Both left and right ovaries are equal in size and are active during the breeding season.
The ovary is oval and flattened laterally with slightly convex lateral and medial surfaces.(31, 76, 87, 96, 105) The free ventral border does not present an ovulation fossa as described in the equine and is convex in young animals and irregular in multiparous females (Figure 5.8).(28, 32, 105) In young females, the ovaries have a relatively smooth surface with only one or two grooves (Figure 5.9).(32) Measurements given for the dromedary ovary vary from 2.6 to 6 cm, 2 to 4 cm and 0.5 to 0.9 cm respectively for length, width, and thickness.(2, 31, 50, 76, 96, 105) The average weight of the ovary ranges from 2 to 15 g.(2, 76) Ovarian weight generally increases with age, ovarian activity, and during pregnancy (corpus-luteum-bearing ovary) (Figure 5.10).(22, 23, 27, 28, 31, 32, 76)
In the Bactrian camel, the ovary has a general morphology and position similar to that described for the dromedary. The proper ligament measures 1.5 to 2.0 cm. The ovary is flat or elliptical in shape. Its size varies from that of a pea to that of a walnut.(26, 51) In the anestrous non-pregnant female, it measures 2.8 to 3.5 cm in length, 2.1 to 3.0 cm in width, and 0.8 to 1.4 cm in thickness.(26)
In the llama, the ovaries are ellipsoid and more globular compared to the alpaca but have a similar general anatomic position to that described for the dromedary. The enclosing ovarian bursa is large. (15, 38, 49, 99) In the llama, the inactive ovaries measure 0.5 x 1.0 x 0.5 cm but nearly double in size when ovarian structures are present (corpora lutea or follicles).(49) The active ovaries measure 1.3 to 2.5 cm x 1 to 2 cm x 0.5 to 1 cm in the llama, and 1.6 ± 0.3 cm x 1.1 ± 0.21 cm x 1.1 ± 0.2 cm in the alpaca. The weight of the ovaries in the alpaca averages 1.87 ± 0.94 g.(13, 38)
Ovarian structures
The ovary is formed by two major parts: the cortex (external part) and the medulla (internal part), measuring respectively 2 to 5 mm and 2 to 4 mm in thickness (Figure 5.11). The organ is surrounded by a tunica albuginea except in the region of the hilus. Follicular activity takes place in the cortex and ovulation can occur anywhere on this surface.
As in other mammalians, the external aspect of the ovary is continuously modified by the type and number of structures present on its surface. These ovarian structures are of 2 types - follicles and corpora lutea - and can be at different stages of development or regression. Contrary to what occurs in most domestic species, there is no cyclic appearance of the corpus luteum in non-bred camelidae females because of the induced nature of ovulation in these species (cf. Physiology). Therefore, in non-bred camelidae females, ovarian activity is mainly follicular and corpora lutea are found only in recently bred or pregnant females.(3, 14, 15, 17, 19, 20, 31, 38, 49, 53, 55, 100, 102, 103, 109)
Follicles
Follicular activity in the camelidae is dominated by 4 types of follicles: small growing follicles, mature pre-ovulatory follicles, regressing follicles, and large anovulatory follicles.(6, 7, 109) Several generations of follicles can be present on the same ovary due to the overlapping of follicular waves in non-bred females (Figure 5.12).
The small growing follicles emerge as a cohort with each new follicular wave. Their number varies from 4 to 8. They are visible on the surface of the ovary as small (2 to 4 mm), slightly-raised vesicles (Figure 5.6).(2, 31, 76, 86, 87, 105, 109) These follicles can been seen alone or in conjunction with other ovarian structures.(2, 31, 76)
The mature pre-ovulatory follicle emerges from the follicular wave after establishing dominance. It measures 10 to 22 mm in size in the dromedary(7, 76, 94, 109) and the Bactrian camel(26) and 7 to 12 mm in the llama and alpaca.(3, 4, 12, 14-16, 19, 100,101) These follicles are spherical and have a thin, clear translucent wall.(2, 19, 31,101) They are turgid and protrude markedly from the ovarian surface.(2, 28, 31, 64, 67, 69, 75, 105) They can be easily detached from the ovarian surface as spherical vesicles (Figure 5.13). The surface of the mature follicle is sometimes highly vascularized.(2, 32) Although some authors have suggested that the size of the mature follicles in the dromedary can be as large as 30 mm,(30-32) our recent observations and those of others suggest that follicles larger than 22 mm should be considered anovulatory because they do not respond to ovulation stimuli.(7, 93, 94, 109)
The appearance of the regressing (atretic) follicles varies according to their stage of regression. At the beginning of regression, the follicular wall becomes opaque and thick (Figure 5.14) and the ovarian diameter regresses slowly until most of the follicle recedes into the ovarian mass. Small atretic follicles are seen on the surface of the follicle as whitish, hard, slightly-raised structures of 1-3 mm, or as dark brown or black spots embedded in the ovarian stroma.(32)
Large anovulatory follicles are present on the ovary in 30 to 50% of non-bred females. Their size and appearance are highly variable and depend on the stage of their evolution (cf. Physiology). They used to be taken for a cystic condition or as large pre-ovulatory follicles.(2, 30, 31) However, recent studies show that these structures are normal in the non-bred females.(7, 94, 109) Anovulatory follicles in the dromedary range in size from 25 to 90 mm. They can have a thin and richly-vascularized or opaque and thick wall (Figure 5.15). The anovulatory follicles contain various amount of serous or hemorrhagic fluid or organized blood clots and fibrin (Figure 5.16 and 5.17). Other follicular structures can be observed on the surface of the ovary or even on the surface of the anovulatory follicle (Figure 5.18).
Similar follicular structures are described in the other members of the camelidae family, especially llamas and alpacas.(5) Follicular dynamics for these species are presented in detail elsewhere in this publication (cf. Female physiology).
Corpus luteum
In the camelidae, the corpus luteum forms after ovulation which occurs 24 to 48 hours after mating. The ovulating follicle becomes increasingly hemorrhagic and collapses at ovulation. The follicular cavity is then filled with blood to form a corpus hemorrhagicum of 10 to 15 mm in diameter. Section of the corpus hemorrhagicum shows a thick luteinized wall with a central blood clot (Figure 5.19).(31) Luteinization of the corpus hemorrhagicum is completed within 4 to 5 days and gives rise to a corpus luteum. The mature corpus luteum is spherical or oblong and protrudes distinctly from the ovarian surface. It has a liver-like consistency. Sagittal section of the corpus luteum shows a reddish brown, richly-vascularized luteal tissue which has a fibrous central zone or sometimes a central cavity of 4 to 6 mm (Figure 5.20).(30-32, 109) In the dromedary, the mature corpus luteum measures 12 to 15 mm in diameter and weighs 1.5 to 2 g.(31, 76)
In the pregnant female, the corpus luteum continues to grow during the first 2 months of gestation, becomes more spherical and protrudes markedly from the ovarian tissue (Figure 5.21).(2, 30-32, 43. 64-66, 76, 87, 91, 105) The corpus luteum of pregnancy has a dark red color with an irregular, highly-vascularized surface. Section of the corpus luteum shows a thin capsule enclosing the luteal tissue. A central fibrous area is usually observed in non-cavitary corpora lutea (Figure 5.22).(30-32)
In the dromedary, size and weight of the corpus luteum of pregnancy are respectively 22.1 ± 6.21 mm (range of 8 to 32) and 4.9 ± 1.31 g (range of 1.6 to 6.2). In our studies, all corpora lutea associated with a pregnancy are at least 20 mm in diameter. In some cases, pregnancy is associated with the presence of 2, 3, or even 4 corpora lutea (Figure 5.23).(30-32, 76, 107) The individual size of these corpora lutea is generally smaller than that of a single corpus luteum of the same pregnancy stage but the overall luteal tissue weight is similar or greater. In the case of double ovulation, one of the corpora lutea can be of normal size while the other is very small.(32)
Regression of the corpus luteum occurs between the 10th and 12th day following a sterile mating.(54, 55, 109) It is characterized by a decrease in size (<12 mm) and vascularization and a change in color, becoming pale or light-brown. The regressing corpus luteum is firm.(30, 31)
Regression of the corpus luteum of pregnancy starts just before parturition and continues during the following few days. The regressing corpora lutea are light brown in color and firm. Their diameter decreases from 16 to 20 mm to 12 to 15 mm. The corpus albicans originating from the complete regression of the corpus luteum of pregnancy is hard, laterally compressed or button shaped, white or gray in color, with no blood vessels on its surface. Corpora albicantia present a dark central zone surrounded by a thick, fibrous capsule with a fibrous central area (Figure 5.24).(30-32)Corpora albicantia of different sizes (5 to 12 mm in diameter) remain on the surface of the ovary of the female for a long time and give the organ its characteristic lobulated appearance, especially in old multiparous females.(30-32)
In the llama and alpaca, corpus luteum evolution and form is similar to that of the dromedary. The size of the mature corpus luteum is 14 to 6 mm.(4, 13, 19, 33-36, 40, 46, 47, 95, 98) The corpus luteum of pregnancy is larger (16 to 18 mm).(4, 12)
In the Bactrian camel, the maximum size of the cyclic corpus luteum (15 mm, range 11 to 23 mm) is attained 7.3 days after copulation. In the absence of pregnancy, the corpus luteum regresses on day 10.6 post-mating. The corpus luteum of pregnancy is larger at 26 mm in diameter on the average (18 to 35 mm). After parturition, the corpus luteum regresses rapidly to 8 to 10 mm then persists for 3 to 4 months.(25, 26)
Histology
Differentiation of ovarian tissue in the dromedary starts at 8 weeks of fetal life. At this stage, the ovary consists mainly of cell clusters or cords tightly packed in the cortex, but loosely arranged in the medulla. It is covered superficially by the germinal epithelium. The primary follicles first appear at 12 weeks and consist of an enlarged primary oocyte surrounded by one follicular cell layer. At 20 to 24 weeks, they either degenerate or continue to develop into a secondary follicle characterized by the presence of more than one cell layer around the oocyte. True follicular cavity is first recognized at 28-32 weeks of gestation. At 32-36 weeks of fetal life, histological and anatomical features of the ovary are similar to those in the prepuberal animal.(41)
Ovarian stroma
The ovarian epithelium is similar to that of cattle. The ovarian stroma is composed of fibrous connective tissue rich in smooth muscle cells. It is divided into two zones: the zona parenchymatosa or cortex, and the zona vasculosa or medulla (Figure 5.25). The connective tissue increases in density from the cortical region to the medulla and hilus.(8) However there is no clear demarcation between the cortex and the medulla. A region of the ovary described as "embryonic remains" is made of a dense connective tissue, presenting projection of the apex of epithelial cells.(8, 76)
Follicular activity is seen in the cortex which contains primordial and secondary follicles as well as tertiary or vesicular follicles. The medulla consists of loose connective tissue containing convoluted, thick-walled ovarian arteries, veins, lymphatics and nerves which enter the ovary at the hilus (Figure 5.26).(32)
A characteristic of the dromedary ovary is the persistence of medullary tubes continuous with mesonephric tubules in the mesovarium. They are lined by epithelium which varies from low cuboidal to columnar. The lumen of these tubes frequently contains secretions. The persistence in the adult camel of medullary tubes in the ovary is linked with persistence of Gartner's duct, since they have the same mesonephric origin.(92)
Ovarian structures
Follicles
Histological features of follicles have been described in the dromedary(2) and the llama.(8)
The general histological organization of follicles is very similar to that of other domestic mammalians. The theca interna and theca externa can only be distinguished from each other in large follicles approaching maturity.(76) The primary follicles are found beneath the tunica albuginea. They appear circular and contain a large ovum surrounded by a single layer of follicular cells. The number of primary follicles varies according to the age of the female (Figure 5.27).(2)
The growing follicles are larger and show a cavity formed by follicular cells and a cumulus oophorus surrounding the ovum. The growing follicle is surrounded by connective tissue (theca folliculi).(2)
The mature follicle protrudes from the surface of the ovary as a large vesicle with a thin wall and a well-defined corona radiata and cumulus oophorus surrounding the ovum.(2) The granulosa cells of large follicles are arranged in columns that radiate from the basement membrane. A rich bed of capillaries is found among the epithelioid cells of the theca interna and the spindle-shaped theca external cells. An area of loose connective tissue separates the theca externa and the dense ovarian stroma (Figure 5.28).(31)
In the llama, the cortex shows many primordial and primary follicles, the majority being primordial. The diameter of primordial follicles and primary follicles ranges respectively from 23.4 to 49.4 µm and from 62.4 to 75.4 µm.(8) This diameter increases 6 to 7 fold (500 µm) in the secondary follicles. The oocyte measures 90 µm in diameter and is larger than that of cattle and sheep. The granulosa is 176 to 200 µm thick and the theca externa measures 44 µm.(8) Tertiary follicles have a diameter of 720 µm. The oocyte in these follicles has a diameter of 112 µm and is surrounded by a zona pellucida of 5.6 µm in thickness and a cumulus oophorus measuring 13-26 µm.(8)
Corpus luteum
The active corpus luteum always appears very vascular and tiny capillaries are seen between the luteal cells (Figure 5.30). The histology of the "cyclic" corpus luteum is similar to that of the pregnancy corpus luteum. They are both covered by a vascular connective tissue capsule which sends septa towards the center.
The luteal tissue is composed mainly of large polygonal granulosa cells supported by a network of reticular tissue. The polygonal luteal cells have a large, spherical and centrally located nucleus containing prominent nucleoli. The cytoplasm is lightly stained and contains fine granules (Figure 5.30).(32, 76) Luteal cells are larger and more numerous in the coipus luteum of pregnancy.(2) Small luteal cells with darkly stained nuclei and scanty cytoplasm are also seen in the luteal tissue. They are considered to be daughter luteal cells of thecal origin (Figure 5.31).(31, 32, 76)
Regressing corpora lutea are characterized by a degeneration of the luteal tissue, a decrease in vascular supply and a progressive invasion by fibrous tissues. The nucleus of luteal cells becomes increasingly pyknotic. All luteal cells decrease in size and disappear gradually to be replaced by a dense fibrous tissue.(76)
In early postpartum, the corpora albicantia are characterized by a degeneration of luteal tissue and its replacement by connective tissue resembling scar tissue. Luteal cells shrink and show increased vacuolization and pyknosis. The blood supply (arterioles) becomes sclerotic.(31, 76, 105) Advanced stages of regression show an increased thickness of the fibrous capsule and trabeculae and fibrosis of the arterioles.(32)
References
1. Abdo. M. S.. H. A. A1 Awkaty, A. S. Janabi, and A. A. A1 Kafawi. 1976. Studies on the endometrium of she-camel (Camelus dromedarius) during the reproductive cycle. Vet. Med. J. Giza. 23:347-357.
2. Abdo, M. S., A. S. Al-Janabi, and A. A. Al-Kafawi. 1969. Studies on the Ovaries of the Female Camel During the Reproductive Cycle and in Conditions Affected with Cysts. Cornell Vet. 59:418-425.
3. Adams, G. P., P. G. Griffin, and O. J. Ginther. 1989. In Situ Morphologic Dynamics of Ovaries, Uterus, and Cervix in Llamas. Biology of Reproduction. 41:551-558.
4. Adams, G. P., J. Sumar, and O. J. Ginther. 1991. Form and function of the corpus luteum in llamas. Animal Reproduction Science. 24:127-138.
5. Adams, G. P., J. Sumar. and O. J. Ginther. 1991. Hemorrhagic Ovarian Follicles in Llamas. Theriogenology. 35:557-568.
6. Anouassi, A. 1984. Activite luteale chez la chamelle non gestante (Camelus dromedarius). These de Doctorat Veterinaire, Institut Agronomique et Veterinaire Hassan II. Rabat, Maroc.
7. Anouassi, A., A. Tibary, M. Adnani, and A. Sghiri. 1994. Preovulatory phase characterization in Camelus dromedarius and induction of ovulation. Proceedings of conference organized by FIS in Niama, Niger.
8. Arias, P., F.-H. Feder, R. Porcel, and J. Cox. 1992. Comparative histological study of the reproductive system of the female llama (Lama guanicoe glama). Anat. Histol. Embryol. 21:314-323.
9. Arthur, G. H., and A. R. El Tigani. 1990. Camel Production in Saudi Arabia. Proceedings of the Workshop 'Is it Possible to Improve the Reproductive Performance of the Camel?' - Paris: 149-160.
10. Barmintsev, Y. N. 1951. Rectal examination of the reproductive organs of the camel. Konevodstvo (Vet. Bull., 22: 725, 1952). 21:38-42.
11. Beddard, F. E. 1902. Mammalia. Cambridge Natural History. 10:285.
12. Bourke, D. A., C. L. Adam, and C. E. Kyle. 1992. Ultrasonography as an aid to controlled breeding in the Llama (Lama glama). Veterinary Record. 130:424-428.
13. Bravo, P.W, and Sumar J. 1984. Some anatomical parameters of the reproductive tract in alpacas. Resum. Invest., San Marcos University, Lima.Peru.
14. Bravo, P. W. 1992. la fase follicular de circlo ovarico y la respuesta de la glandula pituitaria a la copula repetida en la alpaca. Allpak'a. 2.
15. Bravo, P. W. 1994. Reproductive Endocrinology of Llamas and Alpacas. Veterinary Clinics of North America: Food Animal Practice. 10:265-279.
16. Bravo, P. W., and M. E. Fowler. 1986. La aplicacion de la tecnica de ultrasonografia en llamas y alpacas. VII Convencion Intemacional sobre Camelidos Sudamericanos. Potosi, Bolivia. Abstr. 12:12.
17. Bravo, P. W., M. E. Fowler, G. H. Stabenfeldt, and B. L. Lasley. 1990. Ovarian Follicular Dynamics in the Llama. Biology of Reproduction. 43:579-585.
18. Bravo, P. W„ D. Pezo, and V. Alarcon. 1995. Evaluation of early reproductive performance in the postpartum alpaca by progesterone concentrations. Animal Reproduction Science. 39:71-77.
19. Bravo, P. W., and J. Sumar. 1989. Laparascopic Examination of the Ovarian Activity in Alpacas. Animal Reproduction Science. 21:271 -281.
20. Bravo, W., and J. Sumar. 1985. Follicular activity of the ovary in alpacas. In: Proceedings of the International Convention on South American Camelids: 7.
21. Campos, G. K. 1965. Estudio comparative de la imagen citologia vaginal en alpacas con prenez verdadera y pseudo prenez. B.S. Thesis, Fac. Med. Vet., Univ. Nac. Mayor San Marcos, Lima: 1-48.
22. Chahrasbi, A., B. Radmehr, and F. Goulbazhagh. 1975. Anatomy and histology of the reproductive organs of the Iranian camel (Camelus dromedarius). 1. Ovary. J. Vet. Fac. University of Tehran. 30:42-50.
23. Chahrasbi, H., and B. Radmehr. 1975. Anatomical and histological structures of female genitalia in camel (Camelus dromedarius) in Iran. In: Proceedings, XX^' World Veterinary Congress, Thessaloniki, Greece: 41-42.
24. Chaman. E. 1960. Imagen cytologia vaginal y despues de la ovulacion en los alpacos n celo. B.S. Thesis, Fac. Med. Vet. Univ. nacional, San Marcos, Lima
25. Chen. B. X., and Z. X. Yuen. 1984. Pregnancy Diagnosis by Rectal Examination in the Bactrian Camel. In: Cockrill, W.R. (ed.): The Camelid. An all-purpose animal. Scand. Inst. Afric. Studies, Uppsala Sweden.: 399-406.
26. Chen. B. X., and Z. X. Yuen. 1984. Reproductive Pattern of the Bactrian Camel. In: Cockrill,
W.R. (ed.) The Camelid, an all-purpose animal, Scand. Inst. Afric. Studies, Uppsala Sweden: 364-386.
27. Djang, K. T. F., B. A. Harun, J. Kumi-Diaka, H. I. Yusuf, and M. G. Udomah. 1988. Clinical and Anatomical Studies of the Camel (Camelus dromedarius) Genitalia. Theriogenology. 30:1023-1031.
28. El-Wishy. A. B. 1987. Reproduction in the Female Dromedary (Camelus dromedarius): a Review. Animal Reproduction Science. 15:273-297.
29. El-Wishy, A. B. 1988. Reproduction in the Male Dromedary (Camelus dromedarius): a Review. Animal Reproduction Science. 17:217-241.
30. El-Wishy. A. B. 1988. A Study of the genital organs of the female dromedary (Camelus dromedarius). J. Reprod. Fert. 82:587-593.
31. El-Wishy, A. B. 1992. Functional morphology of the ovaries of the dromedary camel. Proc. 1st int.
Camel Conf., Dubai, UAE: 149-154.
32. El-Wishy, A. B., and N. A. Homeida. 1984. Observations on the Ovaries of Slaughtered Camels (Camelus dromedarius). Vet. Med. J. 32:295-313.
33. England. B. G., A. G. Cardozo, and W. C. Foote. 1969. A review of the physiology of reproduction in the New World Camelidae. Intern. Zoo Yearbook. 9:104-109.
34. England, B. G., W. C. Foote, D. H. Matthews, A. G. Cardozo, and S. Riera. 1969. Ovulation and Corpus Luteum Function in the Llama (Lama Glama). J. Endocr. 45:505-513.
35. Femandez-Baca, S. 1970. Luteal Function and the Nature of Reproductive Failures in the Alpaca. Ph.D Thesis, Cornell University, Ithaca, N. Y.
36. Fernandez-Baca, S., W. Hansel, and C. Novoa. 1970. Coipus Luteum Function in the Alpaca. Biology of Reproduction. 3:252-261.
37. Fetaih, H., A. Pospischil, and A. Waldvogel. 1992. Electron Microscopy of the Endometrium of Camels in Normal and some Diseased Conditions. J. Vet. Med A. 39:271-281.
38. Fowler, M. E. 1989. Medicine and Surgery of South American Camelids. Ames, Iowa, Iowa State University Press.
39. Fowler, M. E., and H. J. Olander. 1990. Fetal membranes and ancillary structures of llamas (Lama glama). Am. J. Vet Res. 51:1495-1500. 40. Fuertes, Q. I. 1961. Formacion del cuerpo luteo y algunas observaciones en el endometrio de alpacas jovenes durante los primeros dias de la gestacion. B.S. Thesis, Fac. Med. Vet., Univ. Nac. Mayor San Marcos, Lima: 1-37.
41. George, A. N„ and M. F. A. Fahmy. 1966. Histological study of the developing ovary of the dromedary (Camelus dromedarius). J. Vet. Sci. U.A.R. 3:93-100.
42. Ghazi. S. R., A. Oryan, and H. Pourmirzaei. 1994. Some Aspects of Macroscopic Studies of the Placentation in the Camel (Camelus dromedarius). Anat. Histol. Embryol. 23:337-342.
43. Ghoneim, I. M. 1985. The ovary of the pregnant camel with special reference to corpus luteum. M. V. Sc. Cairo University: 68 pp.
44. Grossman, J. D. 1960. A student’s guide to the anatomy of the camel. Indian Council of agricultural research, New Delhi, India.
45. Hafez, E. S. E. 1955. Foetal-Maternal Attachments in Buffalo and Camel. Indian J. Vet. Sci. 25:109-115.
46. Iniquez, R. L. 1969. Largo de vida del cuerpos luteo y su influencia en al canal reproductivio. B.S. Thesis, Fac. Cienc. Agon., Univ. Mayor San Simon, Cochabamba, Bolivia: 1-54.
47. Iniquez, R. L., and A. Cardozo. 1968. Largo de vida del cuerpos luteo en llamas y su influencia en el canal reproductive. Mem. Segunda Asoc. Latinoam. Prod. Anim., Lima, Pem. 3.
48. Ismail, S. T. 1987. A Review of Reproduction in the Female Camel (Camelus dromedarius). Theriogenology. 28:363-371.
49. Johnson, L. W. 1989. Llama Reproduction. Veterinary Clinics of North America: Food Animal Practice. 5:159-182.
50. Leese, A. S. 1927. A Treatise on the One-humped Camel in health and disease. Hayres & Son Maiden Lane, Stamford, Lincoln Shire.
51. Lesbre, M. F. X. 1906. Recherches Anatomiques Sur les Camelides. Arch. Mus. Nat. VIII.
52. Mahmoud, S. A. 1976. Histological and histochemical studies on the endometrium of the she-camel in relation to ovarian activity. Ph.D. Thesis, Cairo university, Egypt.
53. Marie, M., and A. Anouassi. 1986. Mating-Induced Luteinizing Hormone Surge and Ovulation. Biology of Reproduction. 35:792-798.
54. Marie, M. A. and A. Anouassi. 1987. Induction of luteal activity and progesterone secretion in the nonpregnant one-humped camel (Camelus dromedarius). J. Reprod. Fert. 80:183-192.
55. Marie, M. E. 1987. Bases endocriniennes de la fonction sexuelle chez le dromadaire (Camelus dromedarius). These de doctorat de L'Universite. Paris 6.
56. Merkt, H.. M. Boer, D. Rath, and H. A. Schoon. 1988. The Presence of an Additional Fetal Membrane and its Function in the Newborn Guanaco (Lama guanacoe). Theriogenology. 30:437-439.
57. Meschia, G., H. Prystowsky, A. Hellegers, W. Huckabee, J. Metcalfe, and D. Barron. 1960. Observations on the oxygen supply to the fetal llama. Am. J. Physio. XLV: 284-291.
58. Milne-Edwards, M. H. 1870. Lecon sur la Physiology, 9: 562.
59. Mobarak. A. M., and A. B. El-Wishy. 1971. Uterus of the one-humped camel (Camelus dromedarius) with reference to rectal palpation. Indian J. Anim. Sci. 41:846-855.
60. Morton, W. R. M. 1961. Observations on the full-term foetal membranes of three members of the camelidae (Camelus dromedarius L., Camelus bactrianus L. and Lama Glama L.). J. Anatomy. 25:200-209.
61. Musa, B., and M. E. A. Abusineina. 1976. Pregnancy diagnosis in the camel. Acta. Vet. Beograd. 26:107. 62. Musa, B. E. 1977. A New Epidermal Membrane
Associated with the Foetus of the Camel (Camelus dromedarius). Zbl. Vet. Med. C. Anat. Histol. Embryol. 6:355-358.
63. Musa, B. E. 1979. Reproductive patterns in the female camel (Camelus dromedarius). In: Workshop in Camels, Karthoum, Sudan. Stockholm, Sweden: Inti. Fnd. For Sci.: 279-284.
64. Musa, B. E. 1979. Studies on the Ovary of the Camel (Camelus dromedarius). Sudan J. Vet. Sci. & Anim. Husb. 20:51-56.
65. Musa, B. E., and M. E. Abu Sineina. 1976. Some Observations on Reproduction in the Female Camel (Camelus dromedarius). Acta Veterinaria (Beograd). 26:63-69.
66. Musa, B. E.. and M. E. Abusineina. 1978. Clinical pregnancy diagnosis in the camel and a comparison with bovine pregnancy. The Veterinary Record. 102:7-10.
67. Musa, B. E., and M. E. Abusineina. 1978. The oestrus cycle of the camel (Camelus dromedarius). Veterinary Record. 103:556-557.
68. Musa. B. E., and S. A. Makawi. 1985. Involution of the uterus and the first postpartum heat in the camel (Camelus dromedarius). Conference on Animal Production in Arid Zones, Damascus, Syria, 1985.
69. Nawito, M.. M. R. Shalash. R. Hoppe, and A. M. Rekha. 1967. Reproduction in female camel. Bulletin No. 2, National Research Centre, Cairo.
70. Nawito, M. F. 1967. Some reproductive aspects in the female camel. D. V. M. Thesis Warsaw Agric Univ. Poland: 109 pp.
71. Nayak, R. K. 1977. Scanning Electron Microscopy of the Camel Uterine Tube (Oviduct). Am. J. Vet. Res. 38: 1049-1054.
72. Nosier, M. B. 1973. Histological structure of the mammary glands of the one humped camel (Camelus dromedarius). Indian J. Anim. Sci. 43:639-41.
73. Noumani, A. Z. 1911. The comparative Osteology of the Camel. Journal El Geridah Press.
74. Novoa, C. 1970. REVIEW: Reproduction in the Camelidae. J. Reprod. Fert. 22:3-20.
75. Nur, H. M. 1984. Some Reproductive Aspects and Breeding Patterns of the Somali Camel (Camelus dromedarius). IN: Camel Pastoralism in Somalia. Hussein, M. A. (ed). Proceedings from Workshop in Baydhabo, Mogadishu, Somali Academy of Science and Arts, (Camel forum Working paper, #7): 91-110.
76. Osman, A. 1965. Anatomical Study of the Female Genital System of the One-Humped Camel (Camelus dromedarius) - 1 - The Ovaries. S. J. Vet. Sci. & Anim. Husb. 6:41-52.
77. Osman, A. 1967. Anatomical Study of the Female Genital System of the One-Humped Camel (Camelus dromedarius) - II - The Oviducts. S. J. Vet. Sci. & Anim. Husb. 8:67-77.
78. Osman, A. 1968. Anatomical Study of the Female Genital System of the One-Humped Camel (Camelus dromedarius) - III - The Uterus and Placenta. Sudan J. Vet. Sci. Anim. Husb. 9:477-500.
79. Oumzai. 1996. Contribution a l’etude des modifications histologiques de l'endometre au cours du postpartum chez la femelle du dromadaire (Camelus dromedarius). These de Doctorat Veteinaire de l'lnstitut gronomique et Veterinaire Hassan II, Rabat, Maroc.
80. Owen. 1868. The anatomy of Vertebrates, Vol. II. Fondon, Fongmans, Green and Co..
81. Powers, B. E., L. W. Johnson, L. B. Linton, F. Garry, and J. Smith. 1990. Endometrial biopsy technique and uterine pathologic findings in llamas. JAVMA. 197:1157-1162.
82. Prystowsky. 1960. The placenta and Fetal membranes. Villee (ed.), New York, The Williams and Wilkins Co. p. 159.
83. Savi, P. 1843. Sugli involucri fetali del camelus dromedarius (Fetal envelope of the dromedary camel). In: Miscellanee Medico-Chirurgico, Farmaceutiche, Pisa. Communication of the Session of Zoology and Comparative Anatomy of the Scienziati Italiani, Padua.
84. Sghiri, A. 1988. Evaluation des performances de reproduction d'un troupeau camelin a Laayoune (Camelus dromedarius). These de Doctorat Veterinaire, Insititut Agronomique et Veterinaire Hassan II, Rabat, Maroc.
85. Shalash, M. R 1981. Reproduction in Camels. Egyp. J. Vet. sci. 1118-1152.
86. Shalash, M. R. 1965. Some reproductive aspects in the female camel. World Rev. Anim. Prod. 4:103-108.
87. Shalash. M. R. 1987. Review Article: Reproduction in Camels. Egypt J. Vet. Sci. 24:1 -25.
88. Shalash, M. R.. and M. Nawito. 1964. Some Reproductive Aspects in the Female Camel. 5th ICAR:263-273.
89. Sharina, S. S. 1969. Studies on gestation, birth weight, parturition and involution of uterus in she camel (Camelus dromedarius). Punjab Vet. 8:74-75.
90. Sharnta, S. S., and K. K. Vyas. 1972. Involution of
Uterus and Vulva in Camels. Ceylon Vet. J. 20:9-10.
91. Shawki, H., S. K. Verrna, and R. K. Chandolia. 1992. Studies on Corpus Luteum of Pregnancy in She-Camel (Camelus dromedarius) Throughout the Year. IJAR 13: 80-82.
92. Shehata, R. 1964. Medullary Tubes in the Ovary of the Camel and other Mammals. The Veterinary Record. 76:750-753.
93. Skidmore, J. A., M. Billah, and W. R. Allen. 1995. The ovarian follicular wave pattern in the mated and non-mated dromedary camel (Camelus dromedarius). Journal of Reproduction and Fertility Supplement 49: 545-548
94. Skidmore, J. A.. M. Billah. and W. R. Allen. 1996. The ovarian follicular wave pattern and induction of ovulation in the mated and non mated one humped camel. Journal of Reproduction and Fertility. 106: 186-192.
95. Smith, C. L., A. T. Peter, and D. G. Pugh. 1994. Reproduction in Llamas and Alpacas: A Review. Theriogenology. 41:573-592.
96. Smuts, M. M. S.. and A. J. Bezuidenhout. 1987. Anatomy of the Dromedary. Clarendon Press, Oxford: 230 pp.
97. Stekleniov, E. P. 1968. Des particularites anatomo-morphologiques de la structure et des fonctions physiologiques des trompes de Fallope chez les camelides (genres Lama et Camelus). 6th ICAR. 1:74.
98. Sumar, J. 1983. Studies on Reproductive Pathology in Alpacas. MS Thesis, Dept Obstet and Gynaec., College of Vet Med, Swedish Univ of Agric Sci,. Upsala.
99. Sumar, J. 1985. Reproductive Physiology in South American Camelids. in: Land, R.B., Robinson D.W. (eds), Genetics of Reproduction in Sheep, Butterworths. London: Chapt. 9, 81-95.
100. Sumar, J. 1996. Reproduction in llamas an alpacas. Anim. Reprod. Scien. 42: 405-415
101. Sumar, J., W. Bravo, E. Franco, and et. al. 1985. Laparoscopy in alpacas, llamas and vicunas. In: Proceedings of the International Convention on South American Camelids: 11.
102. Sumar, J., and M. Garcia. 1984. Fisiologia Reproductiva en Los Camelidos Sudamericanos: Alpaca Y Llama. IAEA-SM-292/16:26-27.
103. Sumar, J., and M. Garcia. 1986. Fisiologia de la reproduccion de la alpaca. In: Nuclear and
Related Techniques in Animal Production and Health. IAEA, Vienna: 149-177.
104. Sumar, J., C. Novoa, and S. Femandez-Baca. 1972. Fisiologia Reproductiva Post-Partum en la Alpaca. Rev. Inv. Pec. (IVITA). 1:21-27.
105. Tayeb, M. A. F. 1948. Studies on the Anatomy of the Ovary and Corpus Luteum of the Camel. Brit, vet. J. 104:179-190.
106. Tayeb, M. A. F. 1951. L'appareil genital male du Chameau. Rev. Elevage Med. Vet. Pays Trop. 5: 203-212.
107. Tayeb, M. A. F. 1953. Les organes genitaux de la Chamelle. Rev. Elev. Med. Vet. Pay Trop. 6:17-21.
108. Tayeb, M. F. 1946. The anatomy of the genital organs of the camels, male and female. M. V. Sc. Thesis, Fouad 1st Univ., Cairo, Egypt.
109. Tibary, A., and A. Anouassi. 1996. Ultrasonographyic changes of the reproductive tract in the female camel (Camelus dromedarius) during the follicular cycle and pregnancy. Journal of Camel Practice and Research. 3:71-90
110. Turner. 1875. On the structure of the diffused, the plycotyledonary, and the zonary forms of placenta. J. Anat. Lond. 10:127-177.
111. Van Leenep, E. W. 1964. The placenta of the one humped camel (Camelus dromedarius) during the second half of gestation. Acta Morph. Neesl. Scand. 5:373-379.
112. Vasil'eva, L. P., A. F. Shapilov, and K. Baimukhambetov. 1984. Fetal membranes of the camel. Dokl. Vses. Akad. Skh. Nauk (Vet. Bull., 54: 797, 1984). 5:31-32.
113. Zaganelli, M., and B. C. 1975. Variazioni Metriche Dell'Utero Di Camelus dromedarius. Atti della Societe Italiana Scien Vet. 29:238-240.
Get access to all handy features included in the IVIS website
- Get unlimited access to books, proceedings and journals.
- Get access to a global catalogue of meetings, on-site and online courses, webinars and educational videos.
- Bookmark your favorite articles in My Library for future reading.
- Save future meetings and courses in My Calendar and My e-Learning.
- Ask authors questions and read what others have to say.
Comments (0)
Ask the author
0 comments