Gross and Microscopic Anatomy of the Structures
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GROSS AND MICROSCOPIC ANATOMY OF THE STRUCTURES INVOLVED IN THE PRODUCTION OF SEMINAL FLUID IN THE CHICKEN BY Carl Edward Knight A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE Department of Poultry Science 1967 5// ABSTRACT GROSS AND MICROSCOPIC ANATOMY OF THE STRUCTURES INVOLVED IN THE PRODUCTION OF SEMINAL FLUID IN THE CHICKEN by Carl Edward Knight The gross and microscopic anatomy of the structures involved in the production of seminal fluid in the chicken are discussed. The phallus, round bodies, lymph folds and vascular bodies were found to be the structures directly involved in seminal fluid production. Sur- rounding structures were discussed for orientation and illustrations were presented to complement the discussion. The-cloaca is composed of three chambers; proctodeum, urodeum, and coprodeum. Externally it appears as two lips, a larger dorsal lip and a smaller ventral lip. Its surface is covered with epithelium characteristic of the general body surface. The proctodeal fold arises from the margin of the lips and forms the cloacal opening by its incomplete central fusion. The proctodeal cavity is the most caudal of the cloacal chambers. The phallus, round bodies and lymph folds lie in this chamber. The opening to the cloacal bursa is found in the dorsal aspect of this chamber. The uroproctodeal fold forms the cranial boundary of the proctodeum and the caudal boundary of the urodeum. The urodeum is Carl Edward Knight the middle chamber of the cloaca. The ejaculatory papillae and ureters empty into this chamber. The uroproctodeal fold forms the cranial boundary of the urodeum and the caudal boundary of the coprodeum. The coprodeum is the most cranial of the three chambers of the cloaca and the rectum empties into this chamber. The E, sphincter cloacae, m, contractor cloacae, m. transversus cloacae, m. levator cloacae and m, dialator cloacae are muscles directly associated with the cloaca. Other muscles of the caudal area indirectly act on the cloaca. The a, pudenda interna supplies the vascular bodies and the re- productive structures of the proctodeum. The x, pudenda interna drains the structures and closely parallels the course of the a, pudenda interna. The n, pudenda is closely associated with these vessels. The cloacal ganglion was found to lie close to the vascular body. The vascular bodies lie ventral to the cloaca and are composed of a network of capillaries. Endothelial lined channels were found originating next to the capillaries and draining to larger channels which in turn left the vascular body and entered the phallus, round bodies and lymph folds. The phallus was found to lie in the ventral midline of the procto— deum. Stratified squamous epithelium composes its epithelium. The dermis lies deep to the epidermis with a core of connective tissue lying deep to the dermis. The channels which originate in the vascu- lar body extend into the connective tissue core and the dermis. The round bodies are paired structures which lie on either side of the phallus. Their epidermis is composed of stratified squamous Carl Edward Knight epithelium. Deep to the epidermis is connective tissue which contains the channels. These channels connect with the channels entering the phallus. The lymph folds are paired structures which lie lateral to the round bodies. Their epithelium is simple columnar and contains goblet cells. Deep to the epithelium lies the connective tissue which con- tains the lymph channels. These were found to connect with the phallus and round bodies. In addition numerous lymphocytes were found in the lymph fold. The lymph fold has many of the features of the lower digestive tract but lacks a muscularis mucosa. ACKNOWLEDGEMENTS I would like to thank Dr. A. M. Lucas, Project Leader of the Avian Anatomy project for his assistance, guidance and use of the U. S. Department of Agriculture facilities during the course of this study. I would also like to thank Dr. R. K. Ringer, Professor of Poultry Science and Physiology at Michigan State University for his assistance and constructive criticism in presenting this paper. To Mr. Robert Ewing, illustrator for the Ayian Anatomy project, I owe a special debt of gratitude for his excellent illustrations. His willingness to help and his tremendous talent is greatly appreciated. I would like to thank Mrs. Sandra L. Pangborn, Avian Technologist for the Poultry Science Department and Mrs. Effie M. Dennington, Biologist, U. S. Department of Agriculture for their help in the prepara- tion of slides used in this study and Mrs. Jean A. Fay, Laboratory Technician, U. S. Department of Agriculture for her assistance in the labeling of the drawings. In conclusion, I would like to express my gratitude for the patience and understanding provided by my wife, Janet, during the course of this work. ii TABLE OF CONTENTS Page I ACKNOWLEDGEMENTS . ii II TABLE OF CONTENTS . iii III LIST OF FIGURES . v IV INTRODUCTION . l V REVIEW OF LITERATURE . 3 VI OBJECTIVES . 45 VII PROCEDURE . 46 VIII RESULTS . 48 Gross Description of: Cloaca . 48 Proctodeal Fold . 51 Proctodeal Cavity . 52 Uroproctodeal Fold . 53 Urodeum . 54 Phallus . 54 Round Body . 55 Lymph Fold . 57 Musculature . 58 Arteries . 63 Veins . .‘. 66 vascular Body . 66 Histological Description of: Dorsal Lip . 7O ventral Lip . 73 Proctodeal Fold . 74 Arteries, Veins, Nerves and Muscles Related to the vascular Body . 77 vascular Body . 79 Lymph Channels . 8O Phallus . 82 Round Body . 87 Lymph Fold . 88 iii TABLE OF CONTENTS (continued) Page IX DISCUSSION . 92 X SUMMARY . 96 XI LITERATURE CITED . 99 iv LIST OF FIGURES Figure Page 1. External Surface of Cloaca . 104 2. Cloaca with Dorsal Lip Elevated . 106 3. Cloaca with Proctodeal Fold Pulled Dorsally and Laterally to Expose Proctodeal Cavity . 108 4. Cloaca with Proctodeal Fold and Uroproctodeal Fold Displaced . 110 5. Stereogram of the Cloaca . 112 6. Muscles Related to Cloaca - Caudal View . 114 7. Muscles Related to Cloaca - Lateral View . 116 8. Arterial and Venous Supply to Vascular Body . 118 9. Sagittal Section through Cloaca . 120 10. Frontal Section through Cloaca . 122 11. Low Power of vascular Body Tissue . 124 12. High Power of vascular Body Tissue . 126 13. Sagittal Section through Phallus . 128 14. Three Dimensional Illustration Showing Connections of the Lymph Channels of the vascular Body to Those of the Reproductive Structures of the Proctodeum . 130 INTRODUCTION Sperm are produced in the testis and conveyed to the urodeum through various ducts. When the bird is sexually stimulated, the semen is ejaculated out of the ejaculatory papillae and then flows down a groove which is formed by reproductive structures located in the proc- todeum (the terminal chamber of the cloaca). Just after it leaves the ejaculatory papillae, the semen is diluted by a thin fluid. This thesis is concerned with the structures involved in the production and deliv- erance of this fluid. Conflicting reports and terminology, insufficient information, and the lack of good illustrations of the cloacal area has created some confusion. Previous authors have dealt with various aspects of the anatomy of the cloacal area and the structures involved in dilution of avian semen. Much of the detailed anatomy has been done on birds other than the chicken. The review of literature is presented in some detail to bring terminology used by various authors together. The musculature has been reviewed extensively, as the act of coitus is intimately re- lated to the action of these muscles. Also in studying histological sections, the muscles play an important role in determining the orienta- tion of the tissue. Without this, the sections are very confusing and often times misleading. The arteries, veins, nerves and lymphatics are briefly reviewed. A.more comprehensive study of these structures is needed. 1 2 From the external surface, each item is described as it is en- countered in each chamber of the cloaca. After a description of the cloaca and its contents is presented, the structures related to the cloaca are described including muscles, nerves and blood vessels per- tinent to the area. Following a study of the gross anatomy of the area the histological characteristics, mainly of the reproductive structures, are presented. In reading both the review of literature and the findings of this paper it is helpful to consult the figures presented at the end of this thesis. REVIEW OF LITERATURE Cloaca A. .§£2§§ -- The cloaca, which literally means sewer, is the common receptacle for products of the digestive and urogenital systems. It is composed of three chambers, the proctodeum (Gr. prokto-anus + hodaios - on the way), urodeum (Gr. ouron—urine + hodaios) and coprodeum (Gr. kopros - feces f hodaios) (Stedman, 1961). The avian cloaca is a modification of the crocodilian and saurian types of the terminal end of the digestive and urogenital tracts (Gadow, 1888). In 1829, Barkow described the cloaca of the Gallus_ga11inaceae as having three chambers. The rectum entered the first chamber which he stated was the largest of the three. The middle chamber contained a pair of ureters and a pair of ejaculatory papillae in the male. In the third chamber, he reported an opening to the bursa of Fabricii. In the female, he found the oviduct opening into the middle chamber. Gadow (1888) found that this three—chambered condition existed in the Ratitae, Carinatae and Anatidae which he examined, with the same structures entering the cloaca as reported by Barkow (1829). Gadow referred to these chambers as coprodaeum (coprodeum) which is the first chamber (most cranial chamber) described by Barkow (1829), the urodaeum (urodeum) or middle chamber, and the terminal chamber, the proctodaeum (proctodeum). In this comparative work, Gadow described the difference between the cloacal function of various birds. He indicated that ducks, 3 4 herons and cormorants collected their feces and urine in the coprodeum.