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Gross Appearance of Turkey Cloacae Before and After Single or Multiple Manual Semen Collections
Gross Appearance of Turkey Cloacae Before and After Single or Multiple Manual Semen Collections M. R. BAKST and H. C. CECIL
US Department of Agriculture, ARS, Avian Physiology Laboratory, Beltsville, Maryland 20705 (Received for publication August 13, 1982)
1983 Poultry Science 62:683-689 INTRODUCTION The abdominal massage method of semen collection introduced by Burrows and Quinn (1935, 1937) is the basis of the semen collection techniques used presently with chickens and turkeys. Although modifications of the basic abdominal massage method are employed, the objective of semen collection remains unchanged; that is, to collect high quality semen with a minimum of contamination. In a previous study dealing with the rate of semen replenishment in turkeys (Bakst and Cecil, 1981), red blood cells (RBCs) often contaminated the semen after multiple semen collection (three successive semen collections at 30 to 60 min intervals). Visual inspection of cloacae removed from these males immediately after the last semen collection revealed widespread hemorrhages throughout the proctodeal and urodeal cavities. Therefore, the first objective of this study was to describe the type and extent of injury to the cloacae after single and multiple semen collections. Anatomical descriptions of the turkey male cloacal region are limited to reports by Burrows and Quinn (1937), Komarek (1970), and a description of the poult's cloacal region (for sexing purposes) by Hammond and Marsden (1937). The most comprehensive review of the literature regarding the structure of the avian cloaca, specifically the chicken cloaca, is by Knight (1967, 1970), and more generalized descriptions of the avian cloaca are presented
by King (1975, 1979). Because of the diverse nomenclature and some contradictory observations found in the earlier studies dealing with the anatomy of the turkey cloaca, the second objective of this study was to describe the cloaca of the sexually rested, mature turkey male. When possible, the nomenclature used will be that suggested by King (1979) in Nomina Anatomica Avium. MATERIALS AND METHODS Diamond Hybrid Small White and Nicholas Large White breeder male turkeys were housed separately in floor pens, exposed to a minimum of 14 hr of light per day, and provided free access to feed and water. Males were ejaculated at least once, but generally ejaculated two to three times weekly for at least 6 weeks and then given a 1 week rest before the onset of these studies. All males used averaged at least .2 ml of semen per collection. Controls. Six males sexually rested (no semen collections) for a minimum of 14 days were killed by an intravenous injection of sodium pentobarbital (Somlethal; A. J. Buck & Son, Inc., Cockeysville, MD). Cloacae were excised, examined, and placed in fixative. Group 1. Semen was collected from a total of 19 males either once or at three successive times at 30 to 60 min intervals. With all Group 1 males the semen collector was instructed to perform each cloacal stroke [defined here as
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ABSTRACT The gross appearance of the turkey cloaca was examined before and after single or multiple semen collections. All cloacae exhibited some degree of hemorrhage formation, the extent of which was dependent upon 1) frequency of semen collection, 2) number of cloacal strokes, and 3) individual differences in semen collectors' techniques. Cloacae of males subjected to multiple semen collections of more than four cloacal strokes per semen collection were the most severely injured. Cloacae examined three days after the last semen collection were nearly normal in appearance. It is suggested that cloacal hemorrhages resulting from semen collection can be minimized by 1) reducing the number of cloacal strokes per semen collection to one or two and 2) using the correct hand placement and pressure on the cloaca during the cloacal strokes. (Key words: semen collection, ejaculation, cloaca, turkey, anatomy)
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BAKST AND CECIL
that step during semen collection when sustained hand pressure is applied around the cloacal region for the purpose of expressing semen (see Fig. 10)] with greater and more
..rj^„4..J^„^„,ii...,.r
sustained pressure than would normally be applied. Each male was stroked until semen no longer appeared in the ejaculatory groove, usually in excess of five cloacal strokes. Follow-
__A_,+ S
Downloaded from http://ps.oxfordjournals.org/ at University of Michigan on April 19, 2015
TURKEY CLOACA AFTER SEMEN COLLECTION
RESULTS Controls. The mature turkey male cloaca consisted of three internal compartments: 1) proctodeum, 2) urodeum, and 3) coprodeum (Fig. 1). The proctodeum, the most caudal compartment of the cloaca, contained midventrally the corpora phallica lateralia (phalli) and more craniolaterally, the plicae lymphaticae (lymph folds) (Fig. 2). Together the phalli
and lymph folds constituted the phallus nonprotrudens, the nonintromittent copulatory organ of the turkey, and this will be referred to here as the copulatory apparatus (Fig. 10) (King, 1975). Tumescence of the copulatory apparatus resulted in the formation of the sulcus phalli (ejaculatory groove), a linear depression that separated the turgid folds of the copulatory apparatus (Fig. 10). The dorsal mucosal surface of the proctodeum (dorsal proctodeum) was characterized by a central triangular swelling and longitudinally oriented folds (Knight, 1967) (Fig. 2). The plica uroproctodealis (uroproctodeal fold) merged laterally with the lymph folds and separated the proctodeum from the urodeum (Figs. 1, 3). Within the urodeum were the papillae ductus deferens (papillae) and ostium cloacale ureteris (Fig. 3). Separating the urodeum from the most cranial compartment of the cloaca, the coprodeum, was the plica coprourodealis (coprourodeal fold) (Fig. 1). In fixed specimens the coprodeum was characterized by transversely oriented mucosal folds. A distinct cranial boundary of the coprodeum was absent. However, the junction of the rectum and coprodeum was characterized by an abrupt increase in luminal diameter that continued through the coprourodeal fold. Groups 1 and 2. Immediately After Semen Collection. Cloacae from all Group 1 and Group 2 males subjected to three consecutive semen collections were characterized by primary hemorrhages throughout the cloaca. Hemorrhages were observed between the longitudinal folds of the dorsal proctodeum and, to a lesser degree, on the triangular swelling of the dorsal proctodeum (Figs. 4, 5). Petechiae were on the phallus and lymph folds
FIG. 1. A ventral midline incision through and retraction of the rectum, coprodeum, and phalli reveal the cloacal compartments. The dorsal lip and a short extent of the lateral margins of the rectum and coprodeum were removed to expose the ductus deferens. Petechiae and more expansive hemorrhages are observed on the uroproctodeal fold and to a lesser extent on the dorsal proctodeum after one semen collection (Group 2). The letter abbreviations represent the following structures: P, proctodeum; U, urodeum; C, coprodeum; R, rectum; dd, ductus deferens; up, uroproctodeal fold; cu, coprourodeal fold; If, lymph fold;ts, triangular swelling of the dorsal proctodeum. A millimeter scale is present in most micrographs. FIG. 2. A cloaca from a male killed 8 days after three consecutive semen collections (Group 1). The cloacae (see in Figures 2, 3,4, 5, 8, and 9) were cut at the junction of the dorsal and ventral lips. The dorsal lip was retracted exposing the proctodeum. A midsagittal incision between the two phalli and their subsequent lateral retraction exposed the urodeum. Note that the phalli (p), lymph folds (If), triangular fold (ts), and dorsal proctodeum are unblemished and appear normal. FIG. 3. A cloaca from a male killed 8 days after three consecutive semen collections (Group 2). The papillae (pap), urodeum (U), and uroproctodeal fold are unblemished and appear normal.
Downloaded from http://ps.oxfordjournals.org/ at University of Michigan on April 19, 2015
ing these "exhaustive" semen collections males were killed by cervical dislocation immediately or 1, 3, and 8 days after semen collection. Group 2. Semen from a total of 34 males was also collected once or at three successive times at 30 to 60 min intervals. However, the semen collector was instructed to collect semen in a routine manner. Depending upon the semen collector, each male was stroked two to six times per semen collection. Following these routine semen collections, males were killed immediately or 1, 2, 3, and 8 days after semen collection. Group 2 males were killed by cervical dislocation (20 males) or by an intravenous injection of Somlethal (14 males). Group 3. Five males were subjected to one routine semen collection limited to two cloacal strokes and killed with Somlethal immediately after the semen collection. Group 4. Six males were subjected to one routine semen collection on Monday, Wednesday, and Friday and killed with Somlethal 3 days later (the following Monday). Males in Group 4 were stroked between two and six times. Tissue Preparation. The cloacae of all males were excised and examined visually within 30 min of death. Gross observations were appraised subjectively by the first author.
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Downloaded from http://ps.oxfordjournals.org/ at University of Michigan on April 19, 2015 FIG. 4. Extensive hemorrhage is evident immediately after a single "exhaustive" semen collection (Group 1). Petechiae are abundant on the lymph folds and phalli. FIG. 5. The same cloaca as in Figure 4 after a midsagittal incision in order to expose the urodeum. Note the expanse of the hemorrhage in the urodeum, which includes the papillae (resting at base of the two lower dissecting pins). FIG. 6. Petechiae characterize the luminal surface of a papilla and receptaculum immediately after a single "exhaustive" semen collection (Group 1).
TURKEY CLOACA AFTER SEMEN COLLECTION
uroproctodeal fold, lymph folds, and phalli. One papilla had a single hemorrhage covering its apical half. Group 4 at 3 Days After Semen Collection. Cloacae were characterized by a light brown discoloration around the caudal margin of the dorsal proctodeum and to a lesser extent around the medial face of the lymph folds. Other compartments and structures were comparable to control specimens. DISCUSSION
Our description of cloacae from sexually rested, mature turkey males differs slightly with earlier reports. Komarek (1970) included the copulatory apparatus as part of the urodeum. However, our observations concur with those of Knight (1967) and King (1975, 1979) who described the copulatory apparatus as part of the proctodeum. An apex on each caudoventral surface of the phalli (Burrows and Quinn, 1937) was neither described by Komarek (1970) nor observed in the present study. Conforming to the nomenclature set forth by King (1979) in Nomina Anatomica Avium the correct term for the phalli is the corpus phallica lateralia. There would be no corpus phallicum medianum in the mature turkey male. Both single and multiple semen collection episodes resulted in the formation of primary hemorrhages throughout the proctodeum and urodeum. The number, intensity, and distribution of the hemorrhage sites appeared to vary with the frequency and intensity of the cloacal strokes. For example, forceful and more frequent cloacal strokes with sustained hand pressure on the cloaca (Group 1 males) resulted in more extensive hemorrhage formation than did "routine" semen collection episodes (Groups 2 and 3). Cloacal damage also varied with the semen collectors' techniques. Those individuals using six cloacal strokes per semen collection episode induced greater injury than individuals using two or three cloacal strokes. However, attempts at semen collection from low semen-volume-producing males, regardless of the semen collector, generally resulted in cloacal injury, probably due to both increased hand pressure and frequency of the cloacal strokes. How many cloacal strokes are necessary in order to elicit the maximum volume of high quality semen? Marquez (B. C. Marquez, 1982, personal communication) suggests that generally very little additional semen can be gained after two cloacal strokes.
Downloaded from http://ps.oxfordjournals.org/ at University of Michigan on April 19, 2015
and highly concentrated around the junction of the lymph folds and uroproctodeal fold (Fig. 1). The most severe and extensive hemorrhages were located in the urodeum and included the uroproctodeal fold, urodeal mucosa, and papillae (Fig. 5). The luminal surface of the papillae and distal aspect of the receptaculae also had many petechiae (Fig. 6). Cloacal hemorrhages were less extensive in Group 2 males subjected to a single routine semen collection than in Group 2 males subjected to three consecutive routine semen collections (Fig. 1). In the former group, petechiae generally were limited to the caudal margin of the dorsal proctodeum. Petechiae also characterized the uroproctodeal folds, urodeal mucosa, and papillae (Fig. 9). Groups 1 and 2 at 1, 2, 3, and 8 Days After Semen Collection. One day after the last semen collection the hemorrhagic sites throughout the cloacae of Group 1 males were darker than those in Group 2 males. Both groups had fewer and smaller petechiae on the dorsal proctodeum, uroproctodeal fold, lymph folds, phalli, and papillae (Figs. 7, 8). Cloacae from all Group 1 and 2 males examined 2, 3, and 8 days after the last semen collections showed minor variation in appearance. A light brown discoloration was evident around the caudal margin of the dorsal proctodeum at 2 days (Group 2) and to a lesser extent 3 days after the last semen collection (Groups 1 and 2). Three days after the last semen collection the phalli and lateral surfaces of the lymph folds of all males exhibited no discoloration and were a dull white color. The medial surface of the lymph folds changed from a pinkish color adjacent to the phallus to a light reddish color where it merged with the uroproctodeal fold. This reddish color persisted through the urodeum. Punctate blemishes in the lumina of the papillae and receptaculae only were discernible by microscopy. Except for a slight yellowish discoloration in the dorsal proctodeum and lymph folds, cloacae from Group 1 and 2 males 8 days after the last semen collection were comparable to control specimens. Group 3. Immediately After Semen Collection. Cloacae from Group 3 males subjected to a single semen collection consisting of two cloacal strokes were damaged the least. All cloacae had petechiae between the folds on the caudal margin of the dorsal proctodeum. About half the cloacae had scattered petechiae on the
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Downloaded from http://ps.oxfordjournals.org/ at University of Michigan on April 19, 2015 FIG. 7. Discoloration in the urodeum and between the folds on the dorsal proctodeum 1 day after a single "exhaustive" semen collection is observed (Group 1). FIG. 8. The hemorrhagic appearance persists around the caudal margin of the dorsal proctodeum, urodeum and papillae, one day after three consecutive and "exhaustive" semen collections (Group 1). FIG. 9. Petechiae in the urodeal mucosa are observed with a more diffuse hemorrhage on a papilla immediately after semen collection (Group 2, one semen collection). FIG. 10. Each cloacal stroke should result in tumescence of the copulatory apparatus (seen here) and ejaculation of a semen fraction. Note how the dorsal lip and internally, the dorsal proctodeum, are compressed during a cloacal stroke. (Arrow, ejaculatory groove.)
TURKEY CLOACA AFTER SEMEN COLLECTION T h e dorsal p r o c t o d e u m of all cloacae examined had s o m e petechiae. Such injury m a y b e d u e t o t h e pinching of t h e dorsal p r o c t o d e u m (seen externally as t h e dorsal lip of t h e cloaca) b e t w e e n t h e t h u m b and forefinger during t h e cloacal s t r o k e (Fig. 10). T h e present s t u d y suggests t h a t reducing t h e n u m b e r of cloacal strokes t o o n e or t w o per semen collection minimizes cloacal hemorrhages.
REFERENCES
of spermatozoa from the domestic fowl and turkey. Poultry Sci. 16:19-24. Hammond, J. C , and S. J. Marsden, 1937. Sexing turkeys from hatching to maturity. Poultry Sci. 16:287-288. King, A. S., 1975. Aves urogenital system. Pages 1919—1964 in Sisson and Grossman's The Anatomy of the Domestic Animals. 5th ed. Vol. 2. Ch. 65. R. Getty, ed. W. B. Saunders, Co., Philadelphia, PA. King, A. S., 1979. Systema Urogenitale. Pages 2 8 9 335 in Nomina Anatomica Avium. Julian J. Baumel, ed. Academic Press, New York, NY. Knight, C. E., 1967. Gross and microscopic anatomy of the structures involved in the production of seminal fluid in the chicken. Master's thesis, Michigan State Univ., East Lansing, MI. Knight, C. E., 1970. The anatomy of the structures involved in the erection-dilution mechanism in the male domestic fowl. Ph.D. diss., Michigan State Univ., East Lansing, MI. Komarek, V., 1970. The cloaca of the turkey-cock and of the cock. Acta Vet. Brno. 39:227-234.
NOTE ADDED IN PROOF Recently King (A. S. King, 1 9 8 1 . Phallus. Pages 1 0 7 - 1 4 7 in F o r m and F u n c t i o n in Birds. V o l u m e 2. Ch. 3. A. S. King and J. McLelland, ed. Academic Press, New York, N Y ) described t h e t u r k e y phallus. O u r observations and n o m e n c l a t u r e conform t o t h o s e suggested by King ( 1 9 8 1 ) .
Downloaded from http://ps.oxfordjournals.org/ at University of Michigan on April 19, 2015
Bakst, M. R., and H. C. Cecil, 1981. Changes in the characteristics of turkey ejaculated semen and ductus deferens semen with repeated ejaculations. Reprod. Nutr. Dev. 21:1095-1105. Burrows, W. H., and J. P. Quinn, 1935. A method of obtaining spermatozoa from domestic fowl. Poultry Sci. 14:251-254. Burrows, W. H., and J. P. Quinn, 1937. The collection
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US Department of Agriculture, ARS, Avian Physiology Laboratory, Beltsville, Maryland 20705 (Received for publication August 13, 1982)
1983 Poultry Science 62:683-689 INTRODUCTION The abdominal massage method of semen collection introduced by Burrows and Quinn (1935, 1937) is the basis of the semen collection techniques used presently with chickens and turkeys. Although modifications of the basic abdominal massage method are employed, the objective of semen collection remains unchanged; that is, to collect high quality semen with a minimum of contamination. In a previous study dealing with the rate of semen replenishment in turkeys (Bakst and Cecil, 1981), red blood cells (RBCs) often contaminated the semen after multiple semen collection (three successive semen collections at 30 to 60 min intervals). Visual inspection of cloacae removed from these males immediately after the last semen collection revealed widespread hemorrhages throughout the proctodeal and urodeal cavities. Therefore, the first objective of this study was to describe the type and extent of injury to the cloacae after single and multiple semen collections. Anatomical descriptions of the turkey male cloacal region are limited to reports by Burrows and Quinn (1937), Komarek (1970), and a description of the poult's cloacal region (for sexing purposes) by Hammond and Marsden (1937). The most comprehensive review of the literature regarding the structure of the avian cloaca, specifically the chicken cloaca, is by Knight (1967, 1970), and more generalized descriptions of the avian cloaca are presented
by King (1975, 1979). Because of the diverse nomenclature and some contradictory observations found in the earlier studies dealing with the anatomy of the turkey cloaca, the second objective of this study was to describe the cloaca of the sexually rested, mature turkey male. When possible, the nomenclature used will be that suggested by King (1979) in Nomina Anatomica Avium. MATERIALS AND METHODS Diamond Hybrid Small White and Nicholas Large White breeder male turkeys were housed separately in floor pens, exposed to a minimum of 14 hr of light per day, and provided free access to feed and water. Males were ejaculated at least once, but generally ejaculated two to three times weekly for at least 6 weeks and then given a 1 week rest before the onset of these studies. All males used averaged at least .2 ml of semen per collection. Controls. Six males sexually rested (no semen collections) for a minimum of 14 days were killed by an intravenous injection of sodium pentobarbital (Somlethal; A. J. Buck & Son, Inc., Cockeysville, MD). Cloacae were excised, examined, and placed in fixative. Group 1. Semen was collected from a total of 19 males either once or at three successive times at 30 to 60 min intervals. With all Group 1 males the semen collector was instructed to perform each cloacal stroke [defined here as
683
Downloaded from http://ps.oxfordjournals.org/ at University of Michigan on April 19, 2015
ABSTRACT The gross appearance of the turkey cloaca was examined before and after single or multiple semen collections. All cloacae exhibited some degree of hemorrhage formation, the extent of which was dependent upon 1) frequency of semen collection, 2) number of cloacal strokes, and 3) individual differences in semen collectors' techniques. Cloacae of males subjected to multiple semen collections of more than four cloacal strokes per semen collection were the most severely injured. Cloacae examined three days after the last semen collection were nearly normal in appearance. It is suggested that cloacal hemorrhages resulting from semen collection can be minimized by 1) reducing the number of cloacal strokes per semen collection to one or two and 2) using the correct hand placement and pressure on the cloaca during the cloacal strokes. (Key words: semen collection, ejaculation, cloaca, turkey, anatomy)
684
BAKST AND CECIL
that step during semen collection when sustained hand pressure is applied around the cloacal region for the purpose of expressing semen (see Fig. 10)] with greater and more
..rj^„4..J^„^„,ii...,.r
sustained pressure than would normally be applied. Each male was stroked until semen no longer appeared in the ejaculatory groove, usually in excess of five cloacal strokes. Follow-
__A_,+ S
Downloaded from http://ps.oxfordjournals.org/ at University of Michigan on April 19, 2015
TURKEY CLOACA AFTER SEMEN COLLECTION
RESULTS Controls. The mature turkey male cloaca consisted of three internal compartments: 1) proctodeum, 2) urodeum, and 3) coprodeum (Fig. 1). The proctodeum, the most caudal compartment of the cloaca, contained midventrally the corpora phallica lateralia (phalli) and more craniolaterally, the plicae lymphaticae (lymph folds) (Fig. 2). Together the phalli
and lymph folds constituted the phallus nonprotrudens, the nonintromittent copulatory organ of the turkey, and this will be referred to here as the copulatory apparatus (Fig. 10) (King, 1975). Tumescence of the copulatory apparatus resulted in the formation of the sulcus phalli (ejaculatory groove), a linear depression that separated the turgid folds of the copulatory apparatus (Fig. 10). The dorsal mucosal surface of the proctodeum (dorsal proctodeum) was characterized by a central triangular swelling and longitudinally oriented folds (Knight, 1967) (Fig. 2). The plica uroproctodealis (uroproctodeal fold) merged laterally with the lymph folds and separated the proctodeum from the urodeum (Figs. 1, 3). Within the urodeum were the papillae ductus deferens (papillae) and ostium cloacale ureteris (Fig. 3). Separating the urodeum from the most cranial compartment of the cloaca, the coprodeum, was the plica coprourodealis (coprourodeal fold) (Fig. 1). In fixed specimens the coprodeum was characterized by transversely oriented mucosal folds. A distinct cranial boundary of the coprodeum was absent. However, the junction of the rectum and coprodeum was characterized by an abrupt increase in luminal diameter that continued through the coprourodeal fold. Groups 1 and 2. Immediately After Semen Collection. Cloacae from all Group 1 and Group 2 males subjected to three consecutive semen collections were characterized by primary hemorrhages throughout the cloaca. Hemorrhages were observed between the longitudinal folds of the dorsal proctodeum and, to a lesser degree, on the triangular swelling of the dorsal proctodeum (Figs. 4, 5). Petechiae were on the phallus and lymph folds
FIG. 1. A ventral midline incision through and retraction of the rectum, coprodeum, and phalli reveal the cloacal compartments. The dorsal lip and a short extent of the lateral margins of the rectum and coprodeum were removed to expose the ductus deferens. Petechiae and more expansive hemorrhages are observed on the uroproctodeal fold and to a lesser extent on the dorsal proctodeum after one semen collection (Group 2). The letter abbreviations represent the following structures: P, proctodeum; U, urodeum; C, coprodeum; R, rectum; dd, ductus deferens; up, uroproctodeal fold; cu, coprourodeal fold; If, lymph fold;ts, triangular swelling of the dorsal proctodeum. A millimeter scale is present in most micrographs. FIG. 2. A cloaca from a male killed 8 days after three consecutive semen collections (Group 1). The cloacae (see in Figures 2, 3,4, 5, 8, and 9) were cut at the junction of the dorsal and ventral lips. The dorsal lip was retracted exposing the proctodeum. A midsagittal incision between the two phalli and their subsequent lateral retraction exposed the urodeum. Note that the phalli (p), lymph folds (If), triangular fold (ts), and dorsal proctodeum are unblemished and appear normal. FIG. 3. A cloaca from a male killed 8 days after three consecutive semen collections (Group 2). The papillae (pap), urodeum (U), and uroproctodeal fold are unblemished and appear normal.
Downloaded from http://ps.oxfordjournals.org/ at University of Michigan on April 19, 2015
ing these "exhaustive" semen collections males were killed by cervical dislocation immediately or 1, 3, and 8 days after semen collection. Group 2. Semen from a total of 34 males was also collected once or at three successive times at 30 to 60 min intervals. However, the semen collector was instructed to collect semen in a routine manner. Depending upon the semen collector, each male was stroked two to six times per semen collection. Following these routine semen collections, males were killed immediately or 1, 2, 3, and 8 days after semen collection. Group 2 males were killed by cervical dislocation (20 males) or by an intravenous injection of Somlethal (14 males). Group 3. Five males were subjected to one routine semen collection limited to two cloacal strokes and killed with Somlethal immediately after the semen collection. Group 4. Six males were subjected to one routine semen collection on Monday, Wednesday, and Friday and killed with Somlethal 3 days later (the following Monday). Males in Group 4 were stroked between two and six times. Tissue Preparation. The cloacae of all males were excised and examined visually within 30 min of death. Gross observations were appraised subjectively by the first author.
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Downloaded from http://ps.oxfordjournals.org/ at University of Michigan on April 19, 2015 FIG. 4. Extensive hemorrhage is evident immediately after a single "exhaustive" semen collection (Group 1). Petechiae are abundant on the lymph folds and phalli. FIG. 5. The same cloaca as in Figure 4 after a midsagittal incision in order to expose the urodeum. Note the expanse of the hemorrhage in the urodeum, which includes the papillae (resting at base of the two lower dissecting pins). FIG. 6. Petechiae characterize the luminal surface of a papilla and receptaculum immediately after a single "exhaustive" semen collection (Group 1).
TURKEY CLOACA AFTER SEMEN COLLECTION
uroproctodeal fold, lymph folds, and phalli. One papilla had a single hemorrhage covering its apical half. Group 4 at 3 Days After Semen Collection. Cloacae were characterized by a light brown discoloration around the caudal margin of the dorsal proctodeum and to a lesser extent around the medial face of the lymph folds. Other compartments and structures were comparable to control specimens. DISCUSSION
Our description of cloacae from sexually rested, mature turkey males differs slightly with earlier reports. Komarek (1970) included the copulatory apparatus as part of the urodeum. However, our observations concur with those of Knight (1967) and King (1975, 1979) who described the copulatory apparatus as part of the proctodeum. An apex on each caudoventral surface of the phalli (Burrows and Quinn, 1937) was neither described by Komarek (1970) nor observed in the present study. Conforming to the nomenclature set forth by King (1979) in Nomina Anatomica Avium the correct term for the phalli is the corpus phallica lateralia. There would be no corpus phallicum medianum in the mature turkey male. Both single and multiple semen collection episodes resulted in the formation of primary hemorrhages throughout the proctodeum and urodeum. The number, intensity, and distribution of the hemorrhage sites appeared to vary with the frequency and intensity of the cloacal strokes. For example, forceful and more frequent cloacal strokes with sustained hand pressure on the cloaca (Group 1 males) resulted in more extensive hemorrhage formation than did "routine" semen collection episodes (Groups 2 and 3). Cloacal damage also varied with the semen collectors' techniques. Those individuals using six cloacal strokes per semen collection episode induced greater injury than individuals using two or three cloacal strokes. However, attempts at semen collection from low semen-volume-producing males, regardless of the semen collector, generally resulted in cloacal injury, probably due to both increased hand pressure and frequency of the cloacal strokes. How many cloacal strokes are necessary in order to elicit the maximum volume of high quality semen? Marquez (B. C. Marquez, 1982, personal communication) suggests that generally very little additional semen can be gained after two cloacal strokes.
Downloaded from http://ps.oxfordjournals.org/ at University of Michigan on April 19, 2015
and highly concentrated around the junction of the lymph folds and uroproctodeal fold (Fig. 1). The most severe and extensive hemorrhages were located in the urodeum and included the uroproctodeal fold, urodeal mucosa, and papillae (Fig. 5). The luminal surface of the papillae and distal aspect of the receptaculae also had many petechiae (Fig. 6). Cloacal hemorrhages were less extensive in Group 2 males subjected to a single routine semen collection than in Group 2 males subjected to three consecutive routine semen collections (Fig. 1). In the former group, petechiae generally were limited to the caudal margin of the dorsal proctodeum. Petechiae also characterized the uroproctodeal folds, urodeal mucosa, and papillae (Fig. 9). Groups 1 and 2 at 1, 2, 3, and 8 Days After Semen Collection. One day after the last semen collection the hemorrhagic sites throughout the cloacae of Group 1 males were darker than those in Group 2 males. Both groups had fewer and smaller petechiae on the dorsal proctodeum, uroproctodeal fold, lymph folds, phalli, and papillae (Figs. 7, 8). Cloacae from all Group 1 and 2 males examined 2, 3, and 8 days after the last semen collections showed minor variation in appearance. A light brown discoloration was evident around the caudal margin of the dorsal proctodeum at 2 days (Group 2) and to a lesser extent 3 days after the last semen collection (Groups 1 and 2). Three days after the last semen collection the phalli and lateral surfaces of the lymph folds of all males exhibited no discoloration and were a dull white color. The medial surface of the lymph folds changed from a pinkish color adjacent to the phallus to a light reddish color where it merged with the uroproctodeal fold. This reddish color persisted through the urodeum. Punctate blemishes in the lumina of the papillae and receptaculae only were discernible by microscopy. Except for a slight yellowish discoloration in the dorsal proctodeum and lymph folds, cloacae from Group 1 and 2 males 8 days after the last semen collection were comparable to control specimens. Group 3. Immediately After Semen Collection. Cloacae from Group 3 males subjected to a single semen collection consisting of two cloacal strokes were damaged the least. All cloacae had petechiae between the folds on the caudal margin of the dorsal proctodeum. About half the cloacae had scattered petechiae on the
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BAKST AND CECIL
Downloaded from http://ps.oxfordjournals.org/ at University of Michigan on April 19, 2015 FIG. 7. Discoloration in the urodeum and between the folds on the dorsal proctodeum 1 day after a single "exhaustive" semen collection is observed (Group 1). FIG. 8. The hemorrhagic appearance persists around the caudal margin of the dorsal proctodeum, urodeum and papillae, one day after three consecutive and "exhaustive" semen collections (Group 1). FIG. 9. Petechiae in the urodeal mucosa are observed with a more diffuse hemorrhage on a papilla immediately after semen collection (Group 2, one semen collection). FIG. 10. Each cloacal stroke should result in tumescence of the copulatory apparatus (seen here) and ejaculation of a semen fraction. Note how the dorsal lip and internally, the dorsal proctodeum, are compressed during a cloacal stroke. (Arrow, ejaculatory groove.)
TURKEY CLOACA AFTER SEMEN COLLECTION T h e dorsal p r o c t o d e u m of all cloacae examined had s o m e petechiae. Such injury m a y b e d u e t o t h e pinching of t h e dorsal p r o c t o d e u m (seen externally as t h e dorsal lip of t h e cloaca) b e t w e e n t h e t h u m b and forefinger during t h e cloacal s t r o k e (Fig. 10). T h e present s t u d y suggests t h a t reducing t h e n u m b e r of cloacal strokes t o o n e or t w o per semen collection minimizes cloacal hemorrhages.
REFERENCES
of spermatozoa from the domestic fowl and turkey. Poultry Sci. 16:19-24. Hammond, J. C , and S. J. Marsden, 1937. Sexing turkeys from hatching to maturity. Poultry Sci. 16:287-288. King, A. S., 1975. Aves urogenital system. Pages 1919—1964 in Sisson and Grossman's The Anatomy of the Domestic Animals. 5th ed. Vol. 2. Ch. 65. R. Getty, ed. W. B. Saunders, Co., Philadelphia, PA. King, A. S., 1979. Systema Urogenitale. Pages 2 8 9 335 in Nomina Anatomica Avium. Julian J. Baumel, ed. Academic Press, New York, NY. Knight, C. E., 1967. Gross and microscopic anatomy of the structures involved in the production of seminal fluid in the chicken. Master's thesis, Michigan State Univ., East Lansing, MI. Knight, C. E., 1970. The anatomy of the structures involved in the erection-dilution mechanism in the male domestic fowl. Ph.D. diss., Michigan State Univ., East Lansing, MI. Komarek, V., 1970. The cloaca of the turkey-cock and of the cock. Acta Vet. Brno. 39:227-234.
NOTE ADDED IN PROOF Recently King (A. S. King, 1 9 8 1 . Phallus. Pages 1 0 7 - 1 4 7 in F o r m and F u n c t i o n in Birds. V o l u m e 2. Ch. 3. A. S. King and J. McLelland, ed. Academic Press, New York, N Y ) described t h e t u r k e y phallus. O u r observations and n o m e n c l a t u r e conform t o t h o s e suggested by King ( 1 9 8 1 ) .
Downloaded from http://ps.oxfordjournals.org/ at University of Michigan on April 19, 2015
Bakst, M. R., and H. C. Cecil, 1981. Changes in the characteristics of turkey ejaculated semen and ductus deferens semen with repeated ejaculations. Reprod. Nutr. Dev. 21:1095-1105. Burrows, W. H., and J. P. Quinn, 1935. A method of obtaining spermatozoa from domestic fowl. Poultry Sci. 14:251-254. Burrows, W. H., and J. P. Quinn, 1937. The collection
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