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Intrafollicular Pressure in the Feather Follicle1,2
Intrafollicular Pressure in the Feather Follicle1'2 RONALD A. PETERSON3 AND ROBERT K. RINGER Department
of Poultry Science, Michigan State University, East Lansing, Michigan (Received for publication April 6, 1964)
A
EXPERIMENTAL PROCEDURE
Mature S. C. White Leghorn hens were secured on an operating cradle which held them virtually motionless. A mature feather was randomly selected for studying intrafollicular pressure from either the femoral or caudal feather tract. With procaine used as a local anesthetic, one of the brachial veins was then cannulated for the subsequent injection of sodium pentobarbital. All feather shafts in the immediate area of the chosen feather were cut at the level of the epidermis leaving the selected feather shaft unobstructed. A fine nylon thread was 'Journal Article No. 3304, Michigan Agricultural Experiment Station. 2 'Presented at Poultry Science Assoc. Annual Meeting, Stillwater, Oklahoma, Aug. 19-24, 1963. Report of work done under contract with U.S. Department of Agriculture and authorized by the Research and Marketing Act of 1946. The contract was supervised by the staff of the Poultry Laboratory of the Western Utilization Research & Development Division, Agr. Res. Service, U.S. Dept. of Agr., Albany 10, California. 3 Presented by the senior author in partial fulfillment of the Ph.D. degree.
then stitched superficially into the epithelium around the selected mature feather shaft in a purse string-like manner. The feather shaft was pulled and the empty follicle filled with oil (SAE No. 20). A water-filled polyethylene cannula, connected to a Statham Model P23BB venous transducer, was inserted into the follicle with the purse string being drawn tight and tied. An I. D. .047" X O. D. .067" cannula was used in the retrice feather follicles of the caudal tract while an I. D. .023" X O. D. .038" cannula was employed in measuring intrafollicular pressure in the femoral feather tract. All recordings were made on a Grass Model 5 polygraph. Fifteen (15) to 20 mm. Hg pressure was applied to the system. This initial pressure decreased slightly and then leveled out over a one minute period to a value that was considered as the base pressure in the preanesthetized bird. All subsequent pressures were recorded as a percent of this base. The force necessary to pull a feather from its follicle (feather pulling force) was measured using a spring scale (Klose et at., 1961) throughout the trials and compared to intrafollicular pressure changes. Feathers requiring more than 500 grams of force were considered "tight"; those requiring less than 130 grams were considered "loose." All birds were anesthetized with 3 percent sodium pentobarbital to a depth at which they gave no reflex response when their combs were pinched. RESULTS AND DISCUSSION
Caudal Feather Tract. Intrafollicular pres-
1210
Downloaded from http://ps.oxfordjournals.org/ at Purdue University Libraries ADMN on June 17, 2015
PREVIOUS report by Ostmann, Ringer and Tetzlaff (1963) indicated that anesthetics, sympathetic and parasympathetic blocking agents, and tranquilizing drugs caused the feather to loosen within its follicle. These workers did not clarify the exact role played by the muscles and wall of the feather follicle in feather release. An experiment using intrafollicular pressure as a measurement was designed to study the mechanism involved in the loose and tight states within the feather follicle.
PRESSURE IN FEATHER FOLLICLE
Death Minutes
FIG. 1. The effects of anesthetization and death by cervical dislocation on the intrafollicular pressure of the retrice follicles of the caudal feather tract (11 birds). All percentages given are in relationship to the intrafollicular pressure prior to anesthetization (using 100% as a base). At the peak pressure, the capacity of the recorder was exceeded in a few recordings, thus the break in the solid line.
sure was measured in the number four retrice feather follicle (counting from the end of the row) of the caudal feather tract of 11 birds. The fleshy base of the external tail was secured to a ring support to avoid movement, which would have caused pressure changes within the recording system. Following anesthetization the intrafollicular pressure decreased to 75.8% of the preanesthetized base value (Figure 1). The feather pulling force, measured in the femoral and dorsal feather tracts, decreased until the feathers were loose, (feather pulling force of less than 130 gms.). The feather pulling force decreased more rapidly than did the intrafollicular pressure. After the birds had been anesthetized for 1 to 2
minutes they were killed by cervical dislocation. Thirty seconds after cervical dislocation, intrafollicular pressure decreased to 55.9 percent, while the feather pulling force indicated that the feathers were in a loosened state. About one minute after cervical dislocation, the birds went into mass spasmodic muscle contractions which lasted from 1 to 2 minutes. During this time the feather shafts raised and lowered irregularly over the entire body. When the spasmodic contractions began, the feather pulling force increased as did the intrafollicular pressure which rose to +139.3 percent. Approximately 86 seconds after death the feather pulling force had increased to over 500 gms. The feathers re-
Downloaded from http://ps.oxfordjournals.org/ at Purdue University Libraries ADMN on June 17, 2015
Pre-anes. Anes.
1211
1212
R. A. PETERSON AND R. K. RINGER
Femoral Feather Tract, (a) Death by cervical dislocation. Intrafollicular pressure was measured from feather follicles arbitrarily selected in the femoral feather tract of IS birds. Feather pulling force was measured from feathers arbitrarily selected in the opposite femoral feather tract and compared with intrafollicular pressure through-
160150140 -
S\ Birds killed by cervical dislocation
Feathers tight
Birds killed with sodium Pentobarbital
13o|— 120 110 100 90 80 70 60 50 40 30 20 10 0
X
_L
Death
2
i - IIHHH U
Pre-anes. Anes.
X 3
'42
-112
Minutes FIG. 2. The effects of anesthetization and death by cervical dislocation (IS birds) and death by an overdose of anesthetic (5 birds) on intrafollicular pressure of feather follicles of the femoral feather tract. All percentages given are in relationship to the intrafollicular pressure prior to anesthetization using 100% as a base. At the peak pressure, the capacity of the recorder was exceeded in a few recordings, thus the break in the solid line.
Downloaded from http://ps.oxfordjournals.org/ at Purdue University Libraries ADMN on June 17, 2015
out the trial. Following anesthetization, intrafollicular pressure decreased to 65.2 percent from the 100 percent preanesthetized base pressure (Figure 2), and the feather pulling force decreased until the feathers were loose. The feather pulling force decreased more rapidly than did the intrafollicular pressure. After 1 to 2 minutes in the anesthetized state the birds were killed by cervical dislocation. One minute after death, the intrafollicular pressure had increased to 100 percent, while the feathers remained loose. Approximately one minute after cervical dislocation the birds went
mained in this tightened state, regardless of the intrafollicular pressure level, during the rest of the experiment. When the spasms stopped, intrafollicular pressure decreased to 54.4 percent.
PRESSURE IN FEATHER FOLLICLE
1213
with the increase in feather pulling force were small sporadic pressure changes [similar to those observed in (a)] which continued for approximately 60 minutes after death. Intrafollicular pressure decreased to its lowest level (9.8%) 44 minutes after death with the feathers still remaining in a tightened state. Intrafollicular pressure then increased to 34.1 percent of the preanesthetized base pressure in an average of 103 minutes after death. In birds killed by cervical dislocation, the mass spasmodic muscle contractions possibly initiated a tightened state within the feather follicle much faster than that initiated in birds killed with an overdose of anesthetic. This may be possible because mass muscle spasms have not been observed in birds killed with an overdose of anesthetic. These findings suggest that there are two separate and distinct mechanisms in feather tightening, one occurring prior to death and a second after death.
(b) Death by sodium pentobarbital. In this experiment, the same procedure was used as in (a) except that the birds were killed with an overdose of 3 percent sodium pentobarbital. Death was considered to occur at the time of respiratory failure. When the birds were anesthetized, intrafollicular pressure decreased to 68.7 percent of the 100 percent preanesthetized base pressure (Figure 2). Feather pulling force also decreased to a level at which the feathers were considered to be in the loosened state. After the birds were anesthetized for 1 to 2 minutes they were killed with an overdose of anesthetic. No mass spasmodic muscle contractions were observed, while intrafollicular pressure increased to 73.9 percent. The feather pulling force did not increase until 3.3 minutes after death. The feathers remained tight from this point on regardless of the intrafollicular pressure. Concurrent
SUMMARY Using intrafollicular pressure as a measurement, an experiment was conducted to study the mechanism(s) involved in the loose and tight states within the feather follicle. When birds were anesthetized, intrafollicular pressure decreased, also simultaneously with this the feather pulling force decreased. After death, the feather pulling force and intrafollicular pressure did not appear to be related. Once the feather pulling force had increased to where the feathers were considered in a tightened state after death, it remained there regardless of the intrafollicular pressure. ACKNOWLEDGMENT We are indebted to Mrs. Marjorie J. Tetzlaff, Technician, for assistance rendered.
Downloaded from http://ps.oxfordjournals.org/ at Purdue University Libraries ADMN on June 17, 2015
into mass spasmodic muscle contractions similar to those previously described. Intrafollicular pressure increased to +149.6 percent of the base pressure during this time period. During the spasms which continued for 88 seconds after cervical dislocation, the force necessary to pull the feathers increased until the feathers were tight (500 gms. or over). The feathers remained tight during the remainder of the experiment irrespective of the mtrafollicular pressure level. Intrafollicular pressure decreased to 97.1 percent after the spasms. Small irregular pressure changes continued for 52 minutes after cervical dislocation. Although the intrafollicular pressure fell to 28.4 percent in an average of 39.3 minutes after death, the feathers remained tight. Approximately 120 minutes after death, intrafollicular pressure increased to 68.8 percent. One possible explanation for the increase in intrafollicular pressure might be the onset of rigor in the feather follicle muscles.
1214
R. A. PETERSON AND R. K. RINGER
REFERENCES Klose, A. A., E. P. Mecchi and M. F. Pool, 1961. Observations on factors influencing feather release. Poultry Sci. 40: 1029-1036.
Ostmann, O. W., R. K. Ringer and M. J. Tetzlaff, 1963. The effect of various neuromimetic, anesthetic and tranquilizing drugs on feather release. Poultry Sci. 42: 969-973.
Effects of Protein Level and Source and Grain Source on Performance of Egg Production Stock
(Received for publication April 9, 1964)
T N A typical laying ration cereal grains -*- account for approximately 70% of the total. With the technological advancements made in the grain sorghum industry the substitution of grain sorghum for corn and other cereal grains in laying rations is becoming more evident, especially in the Southwest. However, literature reports of a comparison of the experimental results of substituting grain sorghums for corn and other cereals have been very limited. Adolph and Grau (1956) observed no significant difference in performance between commercial layers on five California poultry ranches for two six-week periods when fed a 17.43% protein predominantly corn ration and an 18.03% protein predominantly milo ration. Berry (1958) compared hens housed in individual cages getting diets with the cereal grain portion (1) all corn, (2) half corn, half milo, and (3) all milo. He reported no significant differences in the performance of the birds receiving these diets. Since the protein levels of the diets were not kept equal, the protein level for the all-milo diet was greater than the all-corn diet which could have been giving the milo diet an advantage over the corn diet. Malik and Quisenberry (1963) used two protein levels, 15 and 18%, with the cereal grain portion consisting of (1) all corn, (2) 75% corn and 25% milo, (3) 50% corn and 50% milo, (4) 25% corn and 75% milo and (5) all
milo. They concluded that birds receiving diets of all-corn, and 50% corn + 50% milo laid significantly more eggs than the other groups. Also birds receiving diets with 100, 75, and 50% of the cereal grain as corn laid significantly larger eggs with a higher feed efficiency than those receiving all milo or 75% of the cereal grain as milo in their diets when the two protein levels were combined. Since the constituents of proteins are amino acids, these amino acids are being continually utilized by the bird either to build new tissues, as in growth and reproduction, or to repair worn tissues. If adequate proteins are lacking in the diet, performance in general will be decreased. Considerable differences are found in the literature concerning the protein level necessary for optimum performance in laying hens. One factor affecting these differences could be that of protein source which ultimately affects the amino acid levels in the diet. The analysis of most feeds for protein involves the chemical determination of the combined nitrogen content. According to Ewing (1951) the nitrogen determined may originate from a number of sources some of which have little or no value as protein. Another factor which could affect the performance of laying hens with reference to protein level and source is that of amino acid imbalance (Naber and Touchburn, 1963). Naber and Touchburn (1963) using
Downloaded from http://ps.oxfordjournals.org/ at Purdue University Libraries ADMN on June 17, 2015
JAMES W. DEATON AND JOHN H. QUISENBERRY Texas A & M University, College Station, Texas
of Poultry Science, Michigan State University, East Lansing, Michigan (Received for publication April 6, 1964)
A
EXPERIMENTAL PROCEDURE
Mature S. C. White Leghorn hens were secured on an operating cradle which held them virtually motionless. A mature feather was randomly selected for studying intrafollicular pressure from either the femoral or caudal feather tract. With procaine used as a local anesthetic, one of the brachial veins was then cannulated for the subsequent injection of sodium pentobarbital. All feather shafts in the immediate area of the chosen feather were cut at the level of the epidermis leaving the selected feather shaft unobstructed. A fine nylon thread was 'Journal Article No. 3304, Michigan Agricultural Experiment Station. 2 'Presented at Poultry Science Assoc. Annual Meeting, Stillwater, Oklahoma, Aug. 19-24, 1963. Report of work done under contract with U.S. Department of Agriculture and authorized by the Research and Marketing Act of 1946. The contract was supervised by the staff of the Poultry Laboratory of the Western Utilization Research & Development Division, Agr. Res. Service, U.S. Dept. of Agr., Albany 10, California. 3 Presented by the senior author in partial fulfillment of the Ph.D. degree.
then stitched superficially into the epithelium around the selected mature feather shaft in a purse string-like manner. The feather shaft was pulled and the empty follicle filled with oil (SAE No. 20). A water-filled polyethylene cannula, connected to a Statham Model P23BB venous transducer, was inserted into the follicle with the purse string being drawn tight and tied. An I. D. .047" X O. D. .067" cannula was used in the retrice feather follicles of the caudal tract while an I. D. .023" X O. D. .038" cannula was employed in measuring intrafollicular pressure in the femoral feather tract. All recordings were made on a Grass Model 5 polygraph. Fifteen (15) to 20 mm. Hg pressure was applied to the system. This initial pressure decreased slightly and then leveled out over a one minute period to a value that was considered as the base pressure in the preanesthetized bird. All subsequent pressures were recorded as a percent of this base. The force necessary to pull a feather from its follicle (feather pulling force) was measured using a spring scale (Klose et at., 1961) throughout the trials and compared to intrafollicular pressure changes. Feathers requiring more than 500 grams of force were considered "tight"; those requiring less than 130 grams were considered "loose." All birds were anesthetized with 3 percent sodium pentobarbital to a depth at which they gave no reflex response when their combs were pinched. RESULTS AND DISCUSSION
Caudal Feather Tract. Intrafollicular pres-
1210
Downloaded from http://ps.oxfordjournals.org/ at Purdue University Libraries ADMN on June 17, 2015
PREVIOUS report by Ostmann, Ringer and Tetzlaff (1963) indicated that anesthetics, sympathetic and parasympathetic blocking agents, and tranquilizing drugs caused the feather to loosen within its follicle. These workers did not clarify the exact role played by the muscles and wall of the feather follicle in feather release. An experiment using intrafollicular pressure as a measurement was designed to study the mechanism involved in the loose and tight states within the feather follicle.
PRESSURE IN FEATHER FOLLICLE
Death Minutes
FIG. 1. The effects of anesthetization and death by cervical dislocation on the intrafollicular pressure of the retrice follicles of the caudal feather tract (11 birds). All percentages given are in relationship to the intrafollicular pressure prior to anesthetization (using 100% as a base). At the peak pressure, the capacity of the recorder was exceeded in a few recordings, thus the break in the solid line.
sure was measured in the number four retrice feather follicle (counting from the end of the row) of the caudal feather tract of 11 birds. The fleshy base of the external tail was secured to a ring support to avoid movement, which would have caused pressure changes within the recording system. Following anesthetization the intrafollicular pressure decreased to 75.8% of the preanesthetized base value (Figure 1). The feather pulling force, measured in the femoral and dorsal feather tracts, decreased until the feathers were loose, (feather pulling force of less than 130 gms.). The feather pulling force decreased more rapidly than did the intrafollicular pressure. After the birds had been anesthetized for 1 to 2
minutes they were killed by cervical dislocation. Thirty seconds after cervical dislocation, intrafollicular pressure decreased to 55.9 percent, while the feather pulling force indicated that the feathers were in a loosened state. About one minute after cervical dislocation, the birds went into mass spasmodic muscle contractions which lasted from 1 to 2 minutes. During this time the feather shafts raised and lowered irregularly over the entire body. When the spasmodic contractions began, the feather pulling force increased as did the intrafollicular pressure which rose to +139.3 percent. Approximately 86 seconds after death the feather pulling force had increased to over 500 gms. The feathers re-
Downloaded from http://ps.oxfordjournals.org/ at Purdue University Libraries ADMN on June 17, 2015
Pre-anes. Anes.
1211
1212
R. A. PETERSON AND R. K. RINGER
Femoral Feather Tract, (a) Death by cervical dislocation. Intrafollicular pressure was measured from feather follicles arbitrarily selected in the femoral feather tract of IS birds. Feather pulling force was measured from feathers arbitrarily selected in the opposite femoral feather tract and compared with intrafollicular pressure through-
160150140 -
S\ Birds killed by cervical dislocation
Feathers tight
Birds killed with sodium Pentobarbital
13o|— 120 110 100 90 80 70 60 50 40 30 20 10 0
X
_L
Death
2
i - IIHHH U
Pre-anes. Anes.
X 3
'42
-112
Minutes FIG. 2. The effects of anesthetization and death by cervical dislocation (IS birds) and death by an overdose of anesthetic (5 birds) on intrafollicular pressure of feather follicles of the femoral feather tract. All percentages given are in relationship to the intrafollicular pressure prior to anesthetization using 100% as a base. At the peak pressure, the capacity of the recorder was exceeded in a few recordings, thus the break in the solid line.
Downloaded from http://ps.oxfordjournals.org/ at Purdue University Libraries ADMN on June 17, 2015
out the trial. Following anesthetization, intrafollicular pressure decreased to 65.2 percent from the 100 percent preanesthetized base pressure (Figure 2), and the feather pulling force decreased until the feathers were loose. The feather pulling force decreased more rapidly than did the intrafollicular pressure. After 1 to 2 minutes in the anesthetized state the birds were killed by cervical dislocation. One minute after death, the intrafollicular pressure had increased to 100 percent, while the feathers remained loose. Approximately one minute after cervical dislocation the birds went
mained in this tightened state, regardless of the intrafollicular pressure level, during the rest of the experiment. When the spasms stopped, intrafollicular pressure decreased to 54.4 percent.
PRESSURE IN FEATHER FOLLICLE
1213
with the increase in feather pulling force were small sporadic pressure changes [similar to those observed in (a)] which continued for approximately 60 minutes after death. Intrafollicular pressure decreased to its lowest level (9.8%) 44 minutes after death with the feathers still remaining in a tightened state. Intrafollicular pressure then increased to 34.1 percent of the preanesthetized base pressure in an average of 103 minutes after death. In birds killed by cervical dislocation, the mass spasmodic muscle contractions possibly initiated a tightened state within the feather follicle much faster than that initiated in birds killed with an overdose of anesthetic. This may be possible because mass muscle spasms have not been observed in birds killed with an overdose of anesthetic. These findings suggest that there are two separate and distinct mechanisms in feather tightening, one occurring prior to death and a second after death.
(b) Death by sodium pentobarbital. In this experiment, the same procedure was used as in (a) except that the birds were killed with an overdose of 3 percent sodium pentobarbital. Death was considered to occur at the time of respiratory failure. When the birds were anesthetized, intrafollicular pressure decreased to 68.7 percent of the 100 percent preanesthetized base pressure (Figure 2). Feather pulling force also decreased to a level at which the feathers were considered to be in the loosened state. After the birds were anesthetized for 1 to 2 minutes they were killed with an overdose of anesthetic. No mass spasmodic muscle contractions were observed, while intrafollicular pressure increased to 73.9 percent. The feather pulling force did not increase until 3.3 minutes after death. The feathers remained tight from this point on regardless of the intrafollicular pressure. Concurrent
SUMMARY Using intrafollicular pressure as a measurement, an experiment was conducted to study the mechanism(s) involved in the loose and tight states within the feather follicle. When birds were anesthetized, intrafollicular pressure decreased, also simultaneously with this the feather pulling force decreased. After death, the feather pulling force and intrafollicular pressure did not appear to be related. Once the feather pulling force had increased to where the feathers were considered in a tightened state after death, it remained there regardless of the intrafollicular pressure. ACKNOWLEDGMENT We are indebted to Mrs. Marjorie J. Tetzlaff, Technician, for assistance rendered.
Downloaded from http://ps.oxfordjournals.org/ at Purdue University Libraries ADMN on June 17, 2015
into mass spasmodic muscle contractions similar to those previously described. Intrafollicular pressure increased to +149.6 percent of the base pressure during this time period. During the spasms which continued for 88 seconds after cervical dislocation, the force necessary to pull the feathers increased until the feathers were tight (500 gms. or over). The feathers remained tight during the remainder of the experiment irrespective of the mtrafollicular pressure level. Intrafollicular pressure decreased to 97.1 percent after the spasms. Small irregular pressure changes continued for 52 minutes after cervical dislocation. Although the intrafollicular pressure fell to 28.4 percent in an average of 39.3 minutes after death, the feathers remained tight. Approximately 120 minutes after death, intrafollicular pressure increased to 68.8 percent. One possible explanation for the increase in intrafollicular pressure might be the onset of rigor in the feather follicle muscles.
1214
R. A. PETERSON AND R. K. RINGER
REFERENCES Klose, A. A., E. P. Mecchi and M. F. Pool, 1961. Observations on factors influencing feather release. Poultry Sci. 40: 1029-1036.
Ostmann, O. W., R. K. Ringer and M. J. Tetzlaff, 1963. The effect of various neuromimetic, anesthetic and tranquilizing drugs on feather release. Poultry Sci. 42: 969-973.
Effects of Protein Level and Source and Grain Source on Performance of Egg Production Stock
(Received for publication April 9, 1964)
T N A typical laying ration cereal grains -*- account for approximately 70% of the total. With the technological advancements made in the grain sorghum industry the substitution of grain sorghum for corn and other cereal grains in laying rations is becoming more evident, especially in the Southwest. However, literature reports of a comparison of the experimental results of substituting grain sorghums for corn and other cereals have been very limited. Adolph and Grau (1956) observed no significant difference in performance between commercial layers on five California poultry ranches for two six-week periods when fed a 17.43% protein predominantly corn ration and an 18.03% protein predominantly milo ration. Berry (1958) compared hens housed in individual cages getting diets with the cereal grain portion (1) all corn, (2) half corn, half milo, and (3) all milo. He reported no significant differences in the performance of the birds receiving these diets. Since the protein levels of the diets were not kept equal, the protein level for the all-milo diet was greater than the all-corn diet which could have been giving the milo diet an advantage over the corn diet. Malik and Quisenberry (1963) used two protein levels, 15 and 18%, with the cereal grain portion consisting of (1) all corn, (2) 75% corn and 25% milo, (3) 50% corn and 50% milo, (4) 25% corn and 75% milo and (5) all
milo. They concluded that birds receiving diets of all-corn, and 50% corn + 50% milo laid significantly more eggs than the other groups. Also birds receiving diets with 100, 75, and 50% of the cereal grain as corn laid significantly larger eggs with a higher feed efficiency than those receiving all milo or 75% of the cereal grain as milo in their diets when the two protein levels were combined. Since the constituents of proteins are amino acids, these amino acids are being continually utilized by the bird either to build new tissues, as in growth and reproduction, or to repair worn tissues. If adequate proteins are lacking in the diet, performance in general will be decreased. Considerable differences are found in the literature concerning the protein level necessary for optimum performance in laying hens. One factor affecting these differences could be that of protein source which ultimately affects the amino acid levels in the diet. The analysis of most feeds for protein involves the chemical determination of the combined nitrogen content. According to Ewing (1951) the nitrogen determined may originate from a number of sources some of which have little or no value as protein. Another factor which could affect the performance of laying hens with reference to protein level and source is that of amino acid imbalance (Naber and Touchburn, 1963). Naber and Touchburn (1963) using
Downloaded from http://ps.oxfordjournals.org/ at Purdue University Libraries ADMN on June 17, 2015
JAMES W. DEATON AND JOHN H. QUISENBERRY Texas A & M University, College Station, Texas