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Radio Telemetric Observations of the Diurnal Changes in Respiration Rate, Heart Rate and Intestinal Motility of Domestic Fowl
SCIENCE STUDIES IN POULTRY BIOLOGY
sylvania State University, University Park, Pennsylvania. Sturkie, P. D., 1965. Avian Physiology. 2nd Edition, Cornell University Press, Ithaca, New York. Talmadge, D. W., 1966. Preparing and Mounting the Chicken Skeleton. 4-H Poultry Study Series. 66-14. University of Connecticut, Storrs, Connecticut. Talmadge, D. W., and W. A. Aho, 1965. Incubation and Embryology of the Chick. Bulletin 65-78, Poultry Science, University of Connecticut, Storrs, Connecticut. Taylor, L. W. Fertility and Hatchability of Chicken and Turkey Eggs. John Wiley & Sons, Inc., New York, New York. Technical Bulletin No. 73, 1953. Social Behavior of the Domestic Fowl. Kansas State University, Manhattan, Kansas. Winter, A. R., and E. M. Funk, 1960. Poultry Science and Practice. 5th Edition. J. B. Lippincott Co., Philadelphia, Penn. Farmer's Bulletin 2216, 1966. Culling hens. U.S. Department of Agriculture.
Radio Telemetric Observations of the Diurnal Changes in Respiration Rate, Heart Rate and Intestinal Motility of Domestic Fowl SHUNZO OSHIMA, KIYOSHI SHIMADA AND TEIICHIRO TONOUE
Department of Animal Physiology, Faculty of Agriculture, Nagoya University, Nagoya, Japan (Received for publication February 13, 1973)
ABSTRACT Patterns of diurnal change in respiration rate, heart rate and cecal motility of broiler chickens were determined by radio-telemetry. Observations were also made on diurnal sleep-wakefulness cycles and eating rhythm. All the observed parameters clearly synchronized with a photoperiod of 14 hours illumination (0500-1900 hrs.) and 10 hours darkness. Respiration rate, heart rate and cecal motility maintained high levels during the light period and decreased sharply on transition from the light to the dark period. The mean values of respiration and heart rates per minute were 35.6 and 335 in the light period and 23.1 and 278 in the dark period. The mean cecal motility, expressed as the number of contractions per hour reached a peak value of 51.6 between 1800 and 1900 hrs., and then decreased sharply to the lowest level of 29.0 after "lights-off." The extent of the appearance of EEG fast waves also decreased to the lowest level at the onset of dark period and increased gradually during the last half of the dark period. POULTRY SCIENCE 53: 503-507, 1974
D
IURNAL activity of domestic fowl is completely dependent on the photoperiod (Foshee et al., 1970). Cain and Wilson (1972) reported that the activity of laying hens was distributed fairly evenly over a 24 hour period under constant illumination. In contrast, deep body temperature in chick-
ens appears to show a clear circadian rhythm even under constant illumination (Winget et al., 1965). In order to observe autonomic functions for a long period, it is essential that the physiological variables are recorded under the unrestrained conditions. Radio-telemetry en-
Downloaded from http://ps.oxfordjournals.org/ at UNIVERSITY OF ARIZONA on June 7, 2015
Jull, M. A., 1952. Poultry Breeding. 3rd Edition. John Wiley & Sons, Inc. New York. Leaflet No. 233, 1958. Selecting Breeding Stock for Broiler Production. U.S. Department of Agriculture, Washington, D.C. Patten, B. M., 1946. Early Embryology of the Chick. 3rd Edition. Blakiston Co., New York, New York. Peterson, R. T., and the Editors of Life, Time, Inc., 1963. The Birds—Life Nature Library. New York. Ridlen, S. F., 1963. The Skeletal Structure of the Chicken. Poultry Project. University of Illinois. Schano, E. A. How to Make A Still Air Incubator. 4-H Poultry Science Project. 6-8-la. New York State College of Ag., Ithaca, New York. Schulman, A. H. Studying Imprinting in Precocial Birds. The Penn-Crest Magazine, Poultry Science Club, Pennsylvania State University, University Park, Pennsylvania. Slucklin, W., 1965. Imprinting and Early Learning. Aldine Publishing Co., Chicago, Illinois. Smith, B. A., 1965. Social Integration in Chicks. The Penn-Crest Magazine, Poultry Science Club, Penn-
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Respiration Rate. One week prior to the experiment a small (0.5 cm. in diameter) pneumatic balloon made of rubber was implanted into the abdominal cavity of the chicken under Nembutal anesthesia (30 mg./kg.). The balloon was located in a position about 6 centimeters ventrolateral to the rectum and connected by polyethylene tubing (I.D. 0.8, O.D., 1.2 mm.) which was led outside body cavity and sutured to the skin. A radio transmitter fabricated in our laboratory (Oshima and Gotoh, 1972) was placed on the back of each bird and connected to the free end of tubing. The transmitted signals were received on an FM radio receiver (Unitec Denshi Corp. Nagoya) and recorded continuously with a pen writing polyrecorder (Toa Electronics, Tokyo). The first one minute tracings were sampled out of every tenminutes recording. The respiration rate per minute was counted in every sample and the data of three birds were averaged for three 24-hr. periods.
MATERIALS AND METHODS
Heart Rate. Rings (4 cm. in diameter) of polyethylene tubing were placed around the base of the wings. The two rings were connected by a piece of plastic plate on which the radio transmitter (Narcobiosystem Corp., Texas) was fixed in place on the back of the bird. A plastic box was used to protect the transmitter from the chickens' pecking. A pair of silver wires (15 cm. in length and 0.5 mm. in diameter) covered with polyethylene tubing, except at the ends, were sutured to the skin of both sides of the breast. The output signals of EKG were fed into an
Male broilers were used. They were raised at a poultry farm until about 6 weeks of age and then placed in individual cages with a photoperiod of 14 hours illumination (05001900 hrs.) and 10 hours darkness. The light intensity was approximately 30 lux at the level of the cage floor. Room temperature fluctuated daily between 16-19° C. Feed and water were supplied ad libitum and a caretaker entered the room once a day between 1000 and 1100 hrs. The outline of the experiment is shown in Table 1.
TABLE 1.—Summary of experimental design
Parameter Respiration rate Heart rate Cecal motility EEG Eating activity
No. of birds 3 3 3 2 3
Body weight (kg.) 1.3-1.5 1.3-1.5 0.8-1.2 1.5-1.6 1.2-1.5
Period recorded with each bird (hrs.) 72 72 72 24 72
Downloaded from http://ps.oxfordjournals.org/ at UNIVERSITY OF ARIZONA on June 7, 2015
ables us to determine the patterning of diurnal changes in the autonomic physiological parameters such as respiration rate, heart rate and intestinal motility being free from the disturbances due to restraint or handling of animals (Mackay, 1968). Sturkie (1963) first used the radio-telemetry in determining heart rate and its diurnal changes in domestic chickens. Cain and Abbott (1970) developed a system for continuous monitoring of the heart rate under unrestrained conditions. Oshima and Sumiya (1971) and Oshima and Gotoh (1972) reported that both respiration rate and cecal motility were successfully monitored over a long period in the chickens. The present study was undertaken to determine the pattern of diurnal changes in heart rate, respiration rate and cecal motility in the chicken. These were then compared with the diurnal sleep-wakef ulness cycles and eating activity patterns which have been observed in our previous experiment.
505
DIURNAL CHANGES
automatic heart rate counter (Unitec Denshi Corp., Nagoya) which selectively counts the R waves of EKG and prints them out. One minute countings of heart rate were taken at ten minutes interval. Each point of the figure represents the mean value of three chickens' recording for three 24-hr. periods.
EEG Recordings. A silver needle type electrode was implanted in the skull in the right hemisphere at a point 3 mm. lateral to midline and 6 mm. anterior to the bregma and at a depth of 1 mm. The reference electrode was placed at a point 5 mm. caudal to the bregma. Both electrodes were secured by dental resin, and a small plastic holder for the transmitter (the same as that used in EKG recordings) was placed over them and fixed with resin. Six to seven days later recording was begun with an EEG recorder (Nihonkoden Co., Tokyo). For analysis one minute tracings were sampled at six-minutes intervals. The duration (in seconds) of fast waves with a frequency higher than 8 Hz. was analyzed in every sample, and the proportion of the time during which the fast waves appeared was calculated. Eating Activity. A device was fabricated in our laboratory. This consisted of a mercury switch placed in a small steel bar set above the feed trough and a pen writing recorder. The steel bar was so set that it moved up and down with the neck movements of birds' pecking. As this device did not give a precise pecking rate, the eating activity was determined by measuring the total time spent by bird in eating per hour. The data obtained was averaged and the ratio of the eating
TIME
OF
DAYIhr)
FIG. 1. Diurnal changes in respiration rate (RR), heart rate (HR), cecal motility (CM), extent of appearance of EEG-fast waves (EEG-FW) and eating activity (EA). Arrows at the top indicate the time of "lights-on" and "off." Each point represents the mean value of the tracings plotted at ten-minute intervals in RR and HR, and six-minute intervals for EEG-FW. Mean values of cecal motility were plotted every ten minutes. The eating activity is expressed as arbitrary unit in which the observed maximum activity (between 0500 to 0600 hrs.) is equal to 1.0. activity/hr. to the maximum eating activity/hr. was calculated over a 24-hr. period. RESULTS EEG sleep-wakefulness cycle, respiration rate, heart rate and cecal motility showed characteristic diurnal changes which synchronized with a photoperiod of 14 hours light and 10 hours dark. All these parameters shifted their levels up and down, corresponding with "lights-on" and "off" (Fig. 1). Eating activity was observed only in the
Downloaded from http://ps.oxfordjournals.org/ at UNIVERSITY OF ARIZONA on June 7, 2015
Cecal Motility. Recording procedure and insertion of the pneumatic balloon was the same as that described in a previous report (Oshima and Sumiya, 1971). The motility was expressed as the number of contractions per 10 minute time period.
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S. OSHIMA, K. SHIMADA AND T. TONOUE
dard errors calculated on the basis of one minute samples or 10 minutes samples at hourly intervals. DISCUSSION Few reports on respiration rate of resting chickens were available previously (Jukes, 1971). The present experiment demonstrated that the respiration rate, heart rate and cecal motility of chickens undergo a clear diurnal rhythm which was dependent on the light and dark cycle. The behavioral eating activity as well as EEG sleep-wakefulness rhythm were also synchronized with the light-dark cycle. However, it should be noted that the minor variation exists in the pattern of diurnal change among respiration rate, heart rate and cecal motility. First, the lowered level after ' 'lights-off' was fairly stable in the heart rate, tended to decrease in respiration rate but tended to increase in heart rate during the dark period. Second, the overshooting in-
TABLE 2.—Respiration rate, heart rate and cecal motility during a 24 hr. period Time of sampling (hr.)' 0100 0200 0300 0400 0500 "lights-on" 0600 0700 0800 0900 1000 1100 1200 1300 1400 1500 1600 1700 1800 1900 "lights-off" 2000 2100 2200 2300 2400
*Mean ± s. e. **number of observations.
Respiration rate (counts/min.) 22.3 ± 1.4*(9)** 23.7 ± 2.2 (9) 19.9 ± 0.8 (8) 20.3 ± 0.7 (7) 32.4 ± 2.2 (9) 36.5 ± 2.0 (6) 33.7 ± 3 . 0 (7) 33.6 ± 2.0 (8) 36.0 ± 1.2 (5) 39.2 ± 1.2 (4) 39.2 ± 1.7 (9) 39.3 ± 2.8 (9) 41.4 ± 2.8 (9) 38.0 ± 2.5 (9) 36.3 ± 1.8 (9) 35.1 ± 1.9 (9) 36.7 ± 1.6 (9) 33.7 ± 2.2 (6) 40.6 ± 2.6 (8) 26.3 ± 1.5 (9) 22.3 ± 1.3 (9) 22.8 ± 1.1 (9) 25.0 ± 2.3 (9) 22.2 ± 1.3 (9)
Heart rate (beats/min.) 279 ± 6.4 (9) 277 ± 4.7 (9) 280 ± 4.7 (9) 277 ± 3.9 (9) 352 ± 8 . 1 (9) 336 ± 4.8 (9) 338 ± 4.8 (9) 340 + 9.1(8) 327 ± 6.5 (9) 347 ± 7.0 (9) 336 ± 6.9 (9) 331 ± 4.2(9) 327 ± 4.8 (9) 326 ± 7.9 (9) 328 ± 4.0 (9) 329 ± 6.0 (9) 343 ± 4 . 1 (9) 341 ± 5.8 (9) 327 ± 3 . 9 (9) 281 ± 2.4 (9) 275 ± 5 . 9 (9) 275 ± 4.4 (9) 275 ± 5 . 7 (9) 283 ± 5 . 8 (9)
Cecal motility (contractions/10 min.) 4.8 ± 0.37 (9) 6.4 ± 0.83 (9) 5.4 ± 0.51 (9) 5.7 ± 0.41 (9) 8.1 ± 0 . 8 4 ( 9 ) 8.1 ± 0 . 7 7 ( 9 ) 6.7 ± 0.41 (9) 6.5 ± 0.64 (9) 6.6 ± 0.77 (9) 9.0 ± 0.91 (7) 7.7 ± 0.72 (9) 8.6 ± 0.82 (9) 7.8 ± 0.69 (9) 7.2 ± 0.81 (9) 7.7 ± 0.73 (9) 7.4 ± 0.72 (9) 7.8 ± 0.72 (9) 8.4 ± 0.86 (9) 6.4 ± 0.40 (9) 4.6 ± 0.26 (9) 5.0 ± 0.33 (9) 5.0 ± 0.46 (9) 5.6 ± 0.29 (9) 5.8 ± 0.29 (9)
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light period. The mean respiration rate was 35.6 per minute in the light period and 23.1 in the dark (30% lower than that of light period). The maximum respiration rate was recorded between 1200 and 1300 hrs. The mean heart rate was 335 in the light period and 278 in the darkness. The highest heart rate was recorded just after the "lightso n " (0500 hrs.). The lowered level of heart rate after "lights-off" remained stable throughout the dark period while the respiration rate tended to decrease in the latter half of the dark period. The high level of cecal motility ranged from 41.8 to 51.6 contractions per hour and reached the peak between 1800 and 1900 hrs., just before "lights-off" and the lowest activity (29.0 contractions per hour) was observed between 1900 and 2000 hrs. just after "lightsoff." This abrupt decrease in cecal motility was followed by a gradual increase during the dark period. In Table 2 are shown the mean respiration rate, heart rate and cecal motility with stan-
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DIURNAL CHANGES
FEBRUARY 14-16.
Further studies are necessary for elucidation of these problems. REFERENCES Bolton, T. B., 1971. Physiology of nervous system. In: D. J. Bell and B. M. Freeman (ed.); Physiology and Biochemistry of the Domestic Fowl, Vol. II, Academic Press, London-New York, pp. 641-673. Cain, J. R., and U. K. Abbott, 1970. A system for diurnal heart rate measurement in chickens. Poultry Sci. 49: 1085-1090. Cain, J. R., and W. O. Wilson, 1972. A test of the circadian rule of Aschoff with chicken hens. J. Interdiscipl. Cycle Res. 3: 77-85. Foshee, D. P., D. M. Centa, C. R. Medaniel and A. Rollo, 1970. Diurnal activity pattern of broilers in a controlled environment. Poultry Sci. 49: 1514— 1518. Jukes, M. G. M., 1971. Control of respiration, In: D. J. Bell and B. M. Freeman (ed.); Physiology and Biochemistry of Domestic Fowl, Vol. I, Academic Press, London-New York, pp. 171-185. Mackay, R. S., 1968. Biomedical Telemetry 1st ed., Wiley & Sons New York-London-Sydney, pp. 1-23. Oshima, S., and T. Sumiya, 1971. A low cost radiopressure transmitter and its application to monitoring cecal motility. J. Physiol. Soc. Japan, 33: 787-788. Oshima, S., and J. Gotoh, 1972. Radio-telemetric studies on the diurnal rhythms of respiratory rate and intestinal motility in chickens. Ann. Rep. Environ. Med. Nagoya Univ. 23: 17-20. Sturkie, P. D., 1963. Heart rate of chickens determined by radio-telemetry during light and dark periods. Poultry Sci. 42: 797-798. Sturkie, P. D., and J. Chilleyzn, 1972. Heart rate changes with age in chickens. Poultry Sci. 51: 906-911. Tummons, J. L., and P. D. Sturkie, 1969. Nervous control of heart rate during excitement in adult White Leghorn cock. Am. J. Physiol. 216: 1437-1440. Watanabe, M., S. Oshima and J. Gotoh, 1973. Influences of restraint and of "external stimuli" on the cecal motility of chickens. Ann. Rep. Environ. Med. Nagoya Univ. 24: 16-19. Winget, C. M., E. G. Averkin and T. B. Fryer, 1965. Quantitative measurement by telemetry of ovulation and oviposition in the fowl. Am. J. Physiol. 209: 853-858.
PACIFIC COAST RENDERERS ASSOCIATION CONVENTION, INN, SCOTTSDALE, ARIZONA.
CAMELBACK
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crease after "lights-on" was observed in the heart rate and cecal motility but not in the respiration rate. These results suggested the existence of not only a major integrating mechanism in controlling the automatic nervous system which is dependent on the light-dark conditions but also the minor specific regulation mechanism responsible for each parameter. Such minor variation in the pattern of diurnal rhythm in automatic nervous functions might be observable only in an experiment in which the birds were not restrained and not excited by handling. Chicken heart rate is considered to be accelerated by the sympathetic nervous system (Sturkie and Chillezyn, 1972) and excited by restraint (Tummons and Sturkie, 1969). Whereas, the cecal motility is significantly inhibited by restraint and this inhibitory effect was thought to be mediated by an adrenergic mechanism (Watanabe et al., 1973). It is generally accepted that the cholinergic and adrenergic mechanisms exert antagonistic effects in the regulation of heart rate and intestinal motility (Bolton, 1971). But, the synchronized decrease in heart rate and cecal motility at the onset of the dark period is hardly explained by a simple sympathetic and parasympathetic regulatory mechanism. If the decrease in heart rate is considered to result from the relative elevation of parasympathetic tonus, it is difficult to explain the decrease in cecal motility, because cecal motility is thought to be accelerated by the parasympathetic nervous system. More complex mechanism must be involved in the autonomic nervous regulation in the diurnal rhythm of cecal motility. Very close correspondence in the pattern of diurnal change in cecal motility and EEG-fast wave might be noted here.
sylvania State University, University Park, Pennsylvania. Sturkie, P. D., 1965. Avian Physiology. 2nd Edition, Cornell University Press, Ithaca, New York. Talmadge, D. W., 1966. Preparing and Mounting the Chicken Skeleton. 4-H Poultry Study Series. 66-14. University of Connecticut, Storrs, Connecticut. Talmadge, D. W., and W. A. Aho, 1965. Incubation and Embryology of the Chick. Bulletin 65-78, Poultry Science, University of Connecticut, Storrs, Connecticut. Taylor, L. W. Fertility and Hatchability of Chicken and Turkey Eggs. John Wiley & Sons, Inc., New York, New York. Technical Bulletin No. 73, 1953. Social Behavior of the Domestic Fowl. Kansas State University, Manhattan, Kansas. Winter, A. R., and E. M. Funk, 1960. Poultry Science and Practice. 5th Edition. J. B. Lippincott Co., Philadelphia, Penn. Farmer's Bulletin 2216, 1966. Culling hens. U.S. Department of Agriculture.
Radio Telemetric Observations of the Diurnal Changes in Respiration Rate, Heart Rate and Intestinal Motility of Domestic Fowl SHUNZO OSHIMA, KIYOSHI SHIMADA AND TEIICHIRO TONOUE
Department of Animal Physiology, Faculty of Agriculture, Nagoya University, Nagoya, Japan (Received for publication February 13, 1973)
ABSTRACT Patterns of diurnal change in respiration rate, heart rate and cecal motility of broiler chickens were determined by radio-telemetry. Observations were also made on diurnal sleep-wakefulness cycles and eating rhythm. All the observed parameters clearly synchronized with a photoperiod of 14 hours illumination (0500-1900 hrs.) and 10 hours darkness. Respiration rate, heart rate and cecal motility maintained high levels during the light period and decreased sharply on transition from the light to the dark period. The mean values of respiration and heart rates per minute were 35.6 and 335 in the light period and 23.1 and 278 in the dark period. The mean cecal motility, expressed as the number of contractions per hour reached a peak value of 51.6 between 1800 and 1900 hrs., and then decreased sharply to the lowest level of 29.0 after "lights-off." The extent of the appearance of EEG fast waves also decreased to the lowest level at the onset of dark period and increased gradually during the last half of the dark period. POULTRY SCIENCE 53: 503-507, 1974
D
IURNAL activity of domestic fowl is completely dependent on the photoperiod (Foshee et al., 1970). Cain and Wilson (1972) reported that the activity of laying hens was distributed fairly evenly over a 24 hour period under constant illumination. In contrast, deep body temperature in chick-
ens appears to show a clear circadian rhythm even under constant illumination (Winget et al., 1965). In order to observe autonomic functions for a long period, it is essential that the physiological variables are recorded under the unrestrained conditions. Radio-telemetry en-
Downloaded from http://ps.oxfordjournals.org/ at UNIVERSITY OF ARIZONA on June 7, 2015
Jull, M. A., 1952. Poultry Breeding. 3rd Edition. John Wiley & Sons, Inc. New York. Leaflet No. 233, 1958. Selecting Breeding Stock for Broiler Production. U.S. Department of Agriculture, Washington, D.C. Patten, B. M., 1946. Early Embryology of the Chick. 3rd Edition. Blakiston Co., New York, New York. Peterson, R. T., and the Editors of Life, Time, Inc., 1963. The Birds—Life Nature Library. New York. Ridlen, S. F., 1963. The Skeletal Structure of the Chicken. Poultry Project. University of Illinois. Schano, E. A. How to Make A Still Air Incubator. 4-H Poultry Science Project. 6-8-la. New York State College of Ag., Ithaca, New York. Schulman, A. H. Studying Imprinting in Precocial Birds. The Penn-Crest Magazine, Poultry Science Club, Pennsylvania State University, University Park, Pennsylvania. Slucklin, W., 1965. Imprinting and Early Learning. Aldine Publishing Co., Chicago, Illinois. Smith, B. A., 1965. Social Integration in Chicks. The Penn-Crest Magazine, Poultry Science Club, Penn-
503
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S. OSHIMA, K. SHIMADA AND T. TONOUE
Respiration Rate. One week prior to the experiment a small (0.5 cm. in diameter) pneumatic balloon made of rubber was implanted into the abdominal cavity of the chicken under Nembutal anesthesia (30 mg./kg.). The balloon was located in a position about 6 centimeters ventrolateral to the rectum and connected by polyethylene tubing (I.D. 0.8, O.D., 1.2 mm.) which was led outside body cavity and sutured to the skin. A radio transmitter fabricated in our laboratory (Oshima and Gotoh, 1972) was placed on the back of each bird and connected to the free end of tubing. The transmitted signals were received on an FM radio receiver (Unitec Denshi Corp. Nagoya) and recorded continuously with a pen writing polyrecorder (Toa Electronics, Tokyo). The first one minute tracings were sampled out of every tenminutes recording. The respiration rate per minute was counted in every sample and the data of three birds were averaged for three 24-hr. periods.
MATERIALS AND METHODS
Heart Rate. Rings (4 cm. in diameter) of polyethylene tubing were placed around the base of the wings. The two rings were connected by a piece of plastic plate on which the radio transmitter (Narcobiosystem Corp., Texas) was fixed in place on the back of the bird. A plastic box was used to protect the transmitter from the chickens' pecking. A pair of silver wires (15 cm. in length and 0.5 mm. in diameter) covered with polyethylene tubing, except at the ends, were sutured to the skin of both sides of the breast. The output signals of EKG were fed into an
Male broilers were used. They were raised at a poultry farm until about 6 weeks of age and then placed in individual cages with a photoperiod of 14 hours illumination (05001900 hrs.) and 10 hours darkness. The light intensity was approximately 30 lux at the level of the cage floor. Room temperature fluctuated daily between 16-19° C. Feed and water were supplied ad libitum and a caretaker entered the room once a day between 1000 and 1100 hrs. The outline of the experiment is shown in Table 1.
TABLE 1.—Summary of experimental design
Parameter Respiration rate Heart rate Cecal motility EEG Eating activity
No. of birds 3 3 3 2 3
Body weight (kg.) 1.3-1.5 1.3-1.5 0.8-1.2 1.5-1.6 1.2-1.5
Period recorded with each bird (hrs.) 72 72 72 24 72
Downloaded from http://ps.oxfordjournals.org/ at UNIVERSITY OF ARIZONA on June 7, 2015
ables us to determine the patterning of diurnal changes in the autonomic physiological parameters such as respiration rate, heart rate and intestinal motility being free from the disturbances due to restraint or handling of animals (Mackay, 1968). Sturkie (1963) first used the radio-telemetry in determining heart rate and its diurnal changes in domestic chickens. Cain and Abbott (1970) developed a system for continuous monitoring of the heart rate under unrestrained conditions. Oshima and Sumiya (1971) and Oshima and Gotoh (1972) reported that both respiration rate and cecal motility were successfully monitored over a long period in the chickens. The present study was undertaken to determine the pattern of diurnal changes in heart rate, respiration rate and cecal motility in the chicken. These were then compared with the diurnal sleep-wakef ulness cycles and eating activity patterns which have been observed in our previous experiment.
505
DIURNAL CHANGES
automatic heart rate counter (Unitec Denshi Corp., Nagoya) which selectively counts the R waves of EKG and prints them out. One minute countings of heart rate were taken at ten minutes interval. Each point of the figure represents the mean value of three chickens' recording for three 24-hr. periods.
EEG Recordings. A silver needle type electrode was implanted in the skull in the right hemisphere at a point 3 mm. lateral to midline and 6 mm. anterior to the bregma and at a depth of 1 mm. The reference electrode was placed at a point 5 mm. caudal to the bregma. Both electrodes were secured by dental resin, and a small plastic holder for the transmitter (the same as that used in EKG recordings) was placed over them and fixed with resin. Six to seven days later recording was begun with an EEG recorder (Nihonkoden Co., Tokyo). For analysis one minute tracings were sampled at six-minutes intervals. The duration (in seconds) of fast waves with a frequency higher than 8 Hz. was analyzed in every sample, and the proportion of the time during which the fast waves appeared was calculated. Eating Activity. A device was fabricated in our laboratory. This consisted of a mercury switch placed in a small steel bar set above the feed trough and a pen writing recorder. The steel bar was so set that it moved up and down with the neck movements of birds' pecking. As this device did not give a precise pecking rate, the eating activity was determined by measuring the total time spent by bird in eating per hour. The data obtained was averaged and the ratio of the eating
TIME
OF
DAYIhr)
FIG. 1. Diurnal changes in respiration rate (RR), heart rate (HR), cecal motility (CM), extent of appearance of EEG-fast waves (EEG-FW) and eating activity (EA). Arrows at the top indicate the time of "lights-on" and "off." Each point represents the mean value of the tracings plotted at ten-minute intervals in RR and HR, and six-minute intervals for EEG-FW. Mean values of cecal motility were plotted every ten minutes. The eating activity is expressed as arbitrary unit in which the observed maximum activity (between 0500 to 0600 hrs.) is equal to 1.0. activity/hr. to the maximum eating activity/hr. was calculated over a 24-hr. period. RESULTS EEG sleep-wakefulness cycle, respiration rate, heart rate and cecal motility showed characteristic diurnal changes which synchronized with a photoperiod of 14 hours light and 10 hours dark. All these parameters shifted their levels up and down, corresponding with "lights-on" and "off" (Fig. 1). Eating activity was observed only in the
Downloaded from http://ps.oxfordjournals.org/ at UNIVERSITY OF ARIZONA on June 7, 2015
Cecal Motility. Recording procedure and insertion of the pneumatic balloon was the same as that described in a previous report (Oshima and Sumiya, 1971). The motility was expressed as the number of contractions per 10 minute time period.
506
S. OSHIMA, K. SHIMADA AND T. TONOUE
dard errors calculated on the basis of one minute samples or 10 minutes samples at hourly intervals. DISCUSSION Few reports on respiration rate of resting chickens were available previously (Jukes, 1971). The present experiment demonstrated that the respiration rate, heart rate and cecal motility of chickens undergo a clear diurnal rhythm which was dependent on the light and dark cycle. The behavioral eating activity as well as EEG sleep-wakefulness rhythm were also synchronized with the light-dark cycle. However, it should be noted that the minor variation exists in the pattern of diurnal change among respiration rate, heart rate and cecal motility. First, the lowered level after ' 'lights-off' was fairly stable in the heart rate, tended to decrease in respiration rate but tended to increase in heart rate during the dark period. Second, the overshooting in-
TABLE 2.—Respiration rate, heart rate and cecal motility during a 24 hr. period Time of sampling (hr.)' 0100 0200 0300 0400 0500 "lights-on" 0600 0700 0800 0900 1000 1100 1200 1300 1400 1500 1600 1700 1800 1900 "lights-off" 2000 2100 2200 2300 2400
*Mean ± s. e. **number of observations.
Respiration rate (counts/min.) 22.3 ± 1.4*(9)** 23.7 ± 2.2 (9) 19.9 ± 0.8 (8) 20.3 ± 0.7 (7) 32.4 ± 2.2 (9) 36.5 ± 2.0 (6) 33.7 ± 3 . 0 (7) 33.6 ± 2.0 (8) 36.0 ± 1.2 (5) 39.2 ± 1.2 (4) 39.2 ± 1.7 (9) 39.3 ± 2.8 (9) 41.4 ± 2.8 (9) 38.0 ± 2.5 (9) 36.3 ± 1.8 (9) 35.1 ± 1.9 (9) 36.7 ± 1.6 (9) 33.7 ± 2.2 (6) 40.6 ± 2.6 (8) 26.3 ± 1.5 (9) 22.3 ± 1.3 (9) 22.8 ± 1.1 (9) 25.0 ± 2.3 (9) 22.2 ± 1.3 (9)
Heart rate (beats/min.) 279 ± 6.4 (9) 277 ± 4.7 (9) 280 ± 4.7 (9) 277 ± 3.9 (9) 352 ± 8 . 1 (9) 336 ± 4.8 (9) 338 ± 4.8 (9) 340 + 9.1(8) 327 ± 6.5 (9) 347 ± 7.0 (9) 336 ± 6.9 (9) 331 ± 4.2(9) 327 ± 4.8 (9) 326 ± 7.9 (9) 328 ± 4.0 (9) 329 ± 6.0 (9) 343 ± 4 . 1 (9) 341 ± 5.8 (9) 327 ± 3 . 9 (9) 281 ± 2.4 (9) 275 ± 5 . 9 (9) 275 ± 4.4 (9) 275 ± 5 . 7 (9) 283 ± 5 . 8 (9)
Cecal motility (contractions/10 min.) 4.8 ± 0.37 (9) 6.4 ± 0.83 (9) 5.4 ± 0.51 (9) 5.7 ± 0.41 (9) 8.1 ± 0 . 8 4 ( 9 ) 8.1 ± 0 . 7 7 ( 9 ) 6.7 ± 0.41 (9) 6.5 ± 0.64 (9) 6.6 ± 0.77 (9) 9.0 ± 0.91 (7) 7.7 ± 0.72 (9) 8.6 ± 0.82 (9) 7.8 ± 0.69 (9) 7.2 ± 0.81 (9) 7.7 ± 0.73 (9) 7.4 ± 0.72 (9) 7.8 ± 0.72 (9) 8.4 ± 0.86 (9) 6.4 ± 0.40 (9) 4.6 ± 0.26 (9) 5.0 ± 0.33 (9) 5.0 ± 0.46 (9) 5.6 ± 0.29 (9) 5.8 ± 0.29 (9)
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light period. The mean respiration rate was 35.6 per minute in the light period and 23.1 in the dark (30% lower than that of light period). The maximum respiration rate was recorded between 1200 and 1300 hrs. The mean heart rate was 335 in the light period and 278 in the darkness. The highest heart rate was recorded just after the "lightso n " (0500 hrs.). The lowered level of heart rate after "lights-off" remained stable throughout the dark period while the respiration rate tended to decrease in the latter half of the dark period. The high level of cecal motility ranged from 41.8 to 51.6 contractions per hour and reached the peak between 1800 and 1900 hrs., just before "lights-off" and the lowest activity (29.0 contractions per hour) was observed between 1900 and 2000 hrs. just after "lightsoff." This abrupt decrease in cecal motility was followed by a gradual increase during the dark period. In Table 2 are shown the mean respiration rate, heart rate and cecal motility with stan-
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FEBRUARY 14-16.
Further studies are necessary for elucidation of these problems. REFERENCES Bolton, T. B., 1971. Physiology of nervous system. In: D. J. Bell and B. M. Freeman (ed.); Physiology and Biochemistry of the Domestic Fowl, Vol. II, Academic Press, London-New York, pp. 641-673. Cain, J. R., and U. K. Abbott, 1970. A system for diurnal heart rate measurement in chickens. Poultry Sci. 49: 1085-1090. Cain, J. R., and W. O. Wilson, 1972. A test of the circadian rule of Aschoff with chicken hens. J. Interdiscipl. Cycle Res. 3: 77-85. Foshee, D. P., D. M. Centa, C. R. Medaniel and A. Rollo, 1970. Diurnal activity pattern of broilers in a controlled environment. Poultry Sci. 49: 1514— 1518. Jukes, M. G. M., 1971. Control of respiration, In: D. J. Bell and B. M. Freeman (ed.); Physiology and Biochemistry of Domestic Fowl, Vol. I, Academic Press, London-New York, pp. 171-185. Mackay, R. S., 1968. Biomedical Telemetry 1st ed., Wiley & Sons New York-London-Sydney, pp. 1-23. Oshima, S., and T. Sumiya, 1971. A low cost radiopressure transmitter and its application to monitoring cecal motility. J. Physiol. Soc. Japan, 33: 787-788. Oshima, S., and J. Gotoh, 1972. Radio-telemetric studies on the diurnal rhythms of respiratory rate and intestinal motility in chickens. Ann. Rep. Environ. Med. Nagoya Univ. 23: 17-20. Sturkie, P. D., 1963. Heart rate of chickens determined by radio-telemetry during light and dark periods. Poultry Sci. 42: 797-798. Sturkie, P. D., and J. Chilleyzn, 1972. Heart rate changes with age in chickens. Poultry Sci. 51: 906-911. Tummons, J. L., and P. D. Sturkie, 1969. Nervous control of heart rate during excitement in adult White Leghorn cock. Am. J. Physiol. 216: 1437-1440. Watanabe, M., S. Oshima and J. Gotoh, 1973. Influences of restraint and of "external stimuli" on the cecal motility of chickens. Ann. Rep. Environ. Med. Nagoya Univ. 24: 16-19. Winget, C. M., E. G. Averkin and T. B. Fryer, 1965. Quantitative measurement by telemetry of ovulation and oviposition in the fowl. Am. J. Physiol. 209: 853-858.
PACIFIC COAST RENDERERS ASSOCIATION CONVENTION, INN, SCOTTSDALE, ARIZONA.
CAMELBACK
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crease after "lights-on" was observed in the heart rate and cecal motility but not in the respiration rate. These results suggested the existence of not only a major integrating mechanism in controlling the automatic nervous system which is dependent on the light-dark conditions but also the minor specific regulation mechanism responsible for each parameter. Such minor variation in the pattern of diurnal rhythm in automatic nervous functions might be observable only in an experiment in which the birds were not restrained and not excited by handling. Chicken heart rate is considered to be accelerated by the sympathetic nervous system (Sturkie and Chillezyn, 1972) and excited by restraint (Tummons and Sturkie, 1969). Whereas, the cecal motility is significantly inhibited by restraint and this inhibitory effect was thought to be mediated by an adrenergic mechanism (Watanabe et al., 1973). It is generally accepted that the cholinergic and adrenergic mechanisms exert antagonistic effects in the regulation of heart rate and intestinal motility (Bolton, 1971). But, the synchronized decrease in heart rate and cecal motility at the onset of the dark period is hardly explained by a simple sympathetic and parasympathetic regulatory mechanism. If the decrease in heart rate is considered to result from the relative elevation of parasympathetic tonus, it is difficult to explain the decrease in cecal motility, because cecal motility is thought to be accelerated by the parasympathetic nervous system. More complex mechanism must be involved in the autonomic nervous regulation in the diurnal rhythm of cecal motility. Very close correspondence in the pattern of diurnal change in cecal motility and EEG-fast wave might be noted here.