Water Metabolism of the Growing Domestic Fowl with Special Reference to Water Balance

BLOOD CONSTITUENTS

Tapper, D. N., and M. R. Kare, 1956. Distribution of glucose in blood of the chicken. Proc. Soc. Exp. Biol. Med. 92: 120-122. Thorbecke, G. J., H. A. Gordon, B. S. Wostmann, M. Wagner and J. A. Reyniers, 19S7. Lymphoid tissue and serum gamma globulin in young germfree chickens. J. Infect. Dis. 101: 237-251.

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Welcker, H., and A. Brandt, 1903. Gewichtswerte der Korperorgane bei dem Menschen und den Tieren. Archiv. f. Anthropologic, 28: 1-89. Yosphe-Purer, Y., J. Fendrich and A. M. Davies, 1953. Estimation of the blood volumes of embryonated hen eggs at different ages. Am. J. Physiol. 175: 178-180.

Water Metabolism of the Growing Domestic Fowl with Special Reference to Water Balance WILLIAM MEDWAY 1 ' 2 AND MORLEY R. KARE

Department of Physiology, New York State Veterinary College, Cornell University, Ithaca, New York

INTRODUCTORY REVIEW

1. Water Intake. The amount of water consumed by a laying White Leghorn hen was reported by Heywang (1941) as approximately 18.2-18.3 gallons per year. He further stated that the amount of water consumed increased with increases in air temperature, live weight and rate of egg production. The converse was also found to be true. It was shown by Kare and Biely (1948) that the addition of increasing amounts of salt to the ration caused a progressive increase in water intake per gram of feed consumed. Increasing the protein content of the feed also increased the water consumption (Wheeler and James, 1950). Patrick (1955), in studying the same relationship, found that increasing the protein content of the feed does not always increase water consumption but may actually decrease it. Heywang (1940), studying the effects of cold drinking water on chick growth and yolk absorption, found that the temperature 1

This is a portion of research completed for the Ph.D. thesis by the senior author. 'Present Address: Department of Medicine, School of Veterinary Medicine, University of Pennsylvania, Philadelphia, Pa.

of the drinking water had no effect on the rate of yolk absorption. Wilson (1948) showed that increasing the environmental temperature caused an increase in water consumption by pullets. He reported that at 95°F. the water consumption was double that at 70°F. Morehouse (1949) measured the water consumption of growing turkeys from hatching to 26 weeks of age. Eley and Hoffmann (1949) showed that feed particle size had no effect on moisture content of the droppings nor on water consumption. Ross et al. (1954) found that water consumption was relatively constant when comparing slow and rapidly growing chicks (2 ml. of water per gram of feed). Jull (1949) reported that a hen in first-year production which lays from 180-240 eggs consumes from 130-180 lb. of water. In other words, it takes approximately 9 lb. of water to produce a dozen eggs. Slinger and Pepper (1955) noticed that when chicks and poults were fed a diet to which penicillin had been added they consumed less water even though they showed the expected growth response. Barott and Pringle (1947) found that chicks on the ninth day after hatching drank approximately 1.6 gm. of water for each gram of

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(Received for publication October 17, 1958)

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W. MEDWAY AND M. R. KARE

In the early literature, the amount of urine voided per day by chickens was reported to be between 500 and 1,000 ml. Other workers have found that volumes of urine excreted were between 86 and 200 ml. (Hester et al., 1940; Hart and Essex, 1942). This discrepancy is probably due to the fact that the measurements were usually made over short periods of time on birds with cannulated ureters or on birds with surgically exteriorized ureters. The effect of various drugs on urine flow has been studied. Weyrauch and Roland (1958) designed an experiment where they sealed off the anus and placed either radiosodium or radioiodine in the cloaca and then used blood for their determination of absorption. It was concluded from these experiments that electrolytes are not reabsorbed from the urine contained in the cloaca. From a physiological basis this would indicate that the role of the cloaca resembles that of the urinary bladder of other mammals. This suggests that the role of the cloaca in water metabolism is a minor one. Dixon (1958) reinvestigated the role played by the cloaca and rectum in absorption. He concluded that little or no water was reabsorbed from ureteral urine by the cloaca or rectum of the hen. The water content of the feces is one of the contributing factors as far as wet litter in poultry houses is concerned. Yushok and Bear (1943) investigating the fertilizing capacity of poultry manure found that freshly voided feces of laying hens contained 77.8 percent water, fresh feces of growing chicks 74.0 percent water, and the fresh feces of two-day-old chicks had 63.1 percent water. No attempt was made by these workers to separate the urine from the feces so that the above results may not be a true indication of fecal water. The feces were also collected on trays and it is difficult to say what precautions were taken to

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feed consumed. Pursuing this study further, Barott and Pringle (1949) found that chicks on the 18th day after hatching drank approximately 1.5S gm. of water for each gram of feed consumed. Humidity had no effect on growth within the range reported. Barott and Pringle (1950) then extended this work to cover birds from 18-32 days of age. The water drunk was approximately 1.75 gm. for every gram of feed consumed. The temperature of the water offered to the chicks in each of the above studies was that of the room. The injection of cockerels with desoxycorticosterone caused an increased water intake, whereas cortisone acetate and ACTH had no effect (Brown et al., 1958a). Adrenalectomy of cockerels caused a decreased water intake (Brown et al., 1958b). 2. The loss of water from the body. In the chicken the cloaca serves as a common excretory pathway for both the urinary and digestive systems. The problem of collecting urine and feces has been overcome by a number of workers who have in essence separated the rectal opening into the cloaca from the ureteral openings by either mechanical or surgical means. Minkowski (1886) circumvented fecal contamination of the urine of the goose by ligating the rectum above the cloaca. Hart and Essex (1942) studied the part played by the cloaca in the water metabolism of the domestic fowl. The ureters in this case were exteriorized. Imabayashi et al. (1955) separated urinary and fecal orifices by performing an "artificial anus" operation. However, their birds did not live longer than three weeks. Dixon and Wilkinson (1957) described a method of exteriorizing the ureters such that the urine and feces could be collected quantitatively in metabolism studies. Their method does not intefere with normal laying. A more detailed review of the development in this area has been compiled by Medway (1958).

WATER METABOLISM AND BALANCE

MATERIALS AND METHODS

Only Single Comb White Leghorn females of a highly inbred strain at Cornell University were used in the experiments. The birds were deprived of feed for 18-24 hours before any determinations were made. Water was supplied ad libitum at all times. The birds were fed commercial rations.

They were housed in appropriate batteries in rooms in which the temperature was kept within the zone of thermal neutrality, this zone being the range of environmental temperature which has little effect on metabolic rate (Dukes, 1955). The relative humidity of the room was not controlled. All determinations where possible were carried out on birds 1, 2, 3, 4, 6, 8, 16, and 32 weeks of age. The fluid intake was obtained by using specially constructed units such that the amount drunk could be measured directly. This unit also had the added feature that a minimum amount of water surface was available for evaporation; however, additional units were used to measure this loss. The water taken in with the feed was easily calculated from feed consumption data and water content of the feed. The excrement was collected in such a way as to limit evaporation of water. This was done quite satisfactorily by collecting the excrement under mineral oil. A special cage was constructed with a wire floor under which stainless steel or enameled pans could be placed. The pans contained mineral oil of sufficient depth for the feces to submerge completely. They were weighed, placed under the birds and the excrement collected for a period of 24 hours. The increase in weight after the collection period represented the total excrement. During this same period the feed consumed, as well as the amount of water drunk, was measured. Before the determinations of intake and outgo were made, the birds were acclimatized to the room and to the feed and water apparatus for at least 24-48 hours. The weights of the birds were recorded at the beginning of each trial. On the first day of the trial, excrement was collected under oil from only half of the experimental units. From the other half of the units the excrement was collected on weighed plastic sheets. This material was then allowed to

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avoid evaporation. Hart and Essex (1942), using birds with artificial anuses reported fecal water values ranging from 65.1 percent to 78.4 percent. Birds with exteriorized ureters showed fecal water values ranging from 74.5 to 93.7 percent. According to Jull (1949), strictly fresh dropping contain approximately 85 percent moisture. Many of the factors which influence the water content of the feces has been reviewed by Medway (1958). Surgical trauma, the injection of desoxycorticosterone, cortisone acetate or ACTH caused an increased water elimination. The elimination of water in each case was probably the result of protein catabolism (Brown et al., 1958a). Medway and Kare (1957) reported evaporative water losses from White Leghorn pullets at various stages of growth. They found that the evaporative water loss was quite high for the first day after hatching. This was followed by a diminution in evaporative water loss for a couple of weeks. It then rose again and leveled off to that of the adult. A study of water metabolism is not complete until one equates the water intake with that lost from the body through all routes. This is a water balance study. Since this information with regard to growing fowl was not available in the literature a need was felt for a study of this type. The effects of various hormonal or environmental stresses on water metabolism could not be appreciated unless balances during normal situations were known.

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W. MEDWAY AND M. R. KAEE TABLE 1.-—Feed and water consumption of growing White Leghorn females

Age (weeks)

No. of birds

1 2 3 4 6 8 16 32

108 96 66 74 50 40 20 20

ml. water Mean chick weight Feed consumpton Water consumption consumed (grams) (gm./chick/day) (ml./chick/day) per gram feed 61.9+ 9.3* 102.8+ 14.7 153.5+ 18.9 250.2+ 32.4 384.4+ 45.6 578.3+ 39.0 1,293.4+138.2 2,035.2 + 199.6

10.4± 1.2 16.9± 2.0 20.5± 1.8 3 2 . 8 ± 4.8 3 8 . 8 ± 7.4 4 9 . 5 ± 6.0 75.5±20.7 136.2±27.9

2.6 2.4 2.6 2.3 2.1 2.4 2.2 3.6

26.6+ 5.2 3 9 . 8 ± 7.6 52.6± 9.4 75.9± 14.5 8 0 . 0 ± 23.7 116.5+ 30.6 165.6± 49.8 483.5+129.9

* Standard deviation.

from the body. In this work, water was studied. The data for the intake which included fluid consumed and water taken in with the feed is shown in Table 2. TABLE 2.—Total daily water intake of White Leghorn females Age

RESULTS Table 1 gives the results of the determination of the daily feed and water consumption by birds during all the ages under investigation. The feed consumption ranged from 10.4 gm./chick/day for the one-weekold bird to 136.2 gm./bird/day for the mature hen. The water consumption showed a parallel increase from 26.6 ml./chick/day for the one-week-old chick to 483.5 ml./day for the mature hen. In investigations involving balance studies, one has to measure both the intake of a particular substance and its output

(weeks)

No. of birds

1 2 3 4 6 8 16 32

108 96 66 74 50 40 20 20

Water intake (ml./bird/day) Liquid

Feed*

26.6 39.8 52.6 75.9 80.0 116.5 165.6 483.5

1.0 1.5 1.9 3.0 3.5 4.5 7.8

Water Total daily wa- intake ter intake ml./gm. (ml./bird) body wt.

12.4

27.5 41.3 54.4 78.9 83.5 121.0 173.4 495.9

.45 .40 .35 .32 .22 .21 .13 .24

* Feed=9.1% moisture.

The total daily water intake for the oneweek-old chick was 27.5 ml. and for the 32-week-old bird it was 495.9 ml. The ratio of water intake per gram of body weight decreased from 0.45 in the one week old to 0.13 in the 16-week-old bird; it then increased to 0.24 in the laying hen.

TABLE 3.—The daily total excrement of growing White Leghorn females Age (weeks)

No. of birds

1 2 3 4 6 8 16 32

108 96 66 74 50 40 20 20

Mean chick weight (gm.)

Total excrement (gm./chick/day)

61.9+ 9.3* 102.8+ 14.7 153.5+ 18.9 250.2+ 32.4 384.4+ 45.6 578.3+ 39.0 1,293.4+138.2 2,035.2 + 199.6

20.6+ 2.4* 28.6+ 4.6 36.2+ 6.3 50.0 + 11.3 56.7 + 15.1 86.2±19.1 122.2 + 44.1 366.7±96.6

Standard deviation.

Total dry feces T o t a l , ex " (gm./chick/day) ™ t p « 4.0 + 0.7 5.6+0.8 7.2 + 0.9 9.0+1.3 10.5 + 4.2 15.0 + 2.4 30.4 + 7.1 47.5 + 8.2

2.0 1.7 1.8 1.5 1.5 1.7 1.6 2.7

% water ^ 80.7 80.4 80.1 81.9 81.6 82.7 75.2 87.0

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dry to constant weight at room temperature. This represented the solid matter of the excrement. The figures obtained were then subtracted from the total excrement of the other half of the units and the resultant figure obtained thus represented the water lost.

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WATER METABOLISM AND BALANCE TABLE 5.—Water balance of growing White Leghorn females

TABLE 4.—Total daily water loss of White Leghorn females Water ou tgo (grams/b rd/day) Age (weeks)

N o . of birds

1 2 3 4 6 8 16 32

108 96 66 74 50 40 20 20

Excrement

Evaporation

16.6 23.0 29.0 41.0 46.2 71.3 91.9 319.2

3.3 6.2 9.2 13.3 19.1 24.8 46.1 53.0

Eggs

34.7

Total daily water loss (gm./bird) 20.0 29.2 38.2 54.2 65.3 96.1 137.9 406.9

Total daily intake (ml./ bird)

Total daily outgo (ml./ bird)

Balance

1 2 3 4 6 8 16 32

61.9 102.8 153.5 250.2 384.4 578.3 1,293.4 2,035.2

27.5 41.3 54.5 78.9 83.5 121.0 173.4 495.9

20.0 29.2 38.2 54.2 65.3 96.1 137.9 406.9

7.6 12.1 16.3 24.6 18.2 24.9 35.5 89.1

Table 3 gives the total daily excrement of the same growing chicks. The water content of the excrement ranged from 75.2-87.0 percent. In the next to the last column of this table is the total excrement per gram of feed. It appears that in the one-week-old chick the efficiency is less than that of the two-week-old bird. This is contrary to what is thought to be the case; further work is needed to establish whether this is true or due to experimental error. The results of the water intake are given in Table 2. From the data it is evident that most of the water intake of the bird comes from the water drunk; this is in contrast THE RELATIONSHIP IN

OF WATER INTAKE AND WATER LOSS AGAINST BOOT WEIGHT

GROWING

FEMALE

WHITE

LEGHORN

CHICKENS

DISCUSSION In Table 1 the data for the feed and water consumption per day by the chicks of the various ages are given; the last column of the table shows the ratio of milliliters water consumed per gram of feed.

/ ^ 200

400

600

800 1000 1200 MOO CHICKEN WEIGHT (GM)

FIG.

1

1600

«00

2000

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Table 3 gives the results of the experiments dealing with the total excrement of the birds. The total excrement for the oneweek-old bird was 20.6 gm./day, this increased steadily; for the mature laying hen the total excrement was 366.7 gm./day. The total dry feces was 4.0 gm./day for the one-week-old chick and 47.5 gm./day for the 32-week-old hen. The total excrement per gram of feed consumed ranged from 1.5 to 2.7. The amount of water present in the total excrement as shown in the last column ranged from 75.2 to 87.0 percent. The total water loss of the birds, that is, through evaporation as well as in the excrement, is shown in Table 4. The total daily water loss through all routes was 20.0 gm./ day for the one-week-old chick and 406.9 gm./day for the 32-week-old bird. Table 5 shows the results of the balance studies. It can be seen that the birds were in a positive balance throughout; this was expected since the birds were growing throughout the period of study. The water balance is shown graphically in Figure 1. The ruled area is the difference between the water intake and the water loss.

Age (weeks)

Mean chick weight (gm.)

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W. MEDWAY AND M. R. KARE

has been shown that during the period of a half to one hour after handling the birds the urine flow is greatly increased. Changes in environment probably also play a large role in determining the volume of urine voided per day. In experiments where the rectum has been plugged with cotton or the ureters cannulated an increased flow was observed. This increase was probably due to handling and anesthesia of the bird. In the studies being reported here no attempt was made to separate the urine from the feces or to determine fluid lost through the oviduct of the laying hen. In all ages studied the birds were in a positive balance which was the expected result since the birds were all normal and growing (Fig. 1). The amount of water retained in the body by the young chick was relatively higher proportionately than that in the laying hen of 32 weeks of age. CONCLUSIONS The milliliters of water drunk per gram of feed consumed ranged from 2.1-2.6 for all ages studied with the exception of the 32-week-old laying hens; here the amount was 3.6 ml./gm. of feed. This increase in water consumption was believed to be due to the water lost in the egg. The total excrement per gram of feed consumed ranged from 1.5-2.0 gm. Here the 32-week-old laying bird voided 2.7 grams of excrement per gram of feed consumed. The daily water intake in liquid and that present in the feed was 27.5 ml. per bird for the one-week-old chick and 495.9 ml. per bird for the 32-week-old hen. The total daily water loss was about 20.0 gm./bird for the one-week-old chick and 406.9 gm./bird for the 32-week-old laying hen. The birds were found to be in a positive water balance throughout the studies. This was probably related to growth.

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with other animals that eat a very succulent diet. The one-week-old chick consumes on the average 27.5 ml. of water per day; this increases steadily while the chick grows till finally at 32 weeks of age the water intake is about 495.9 ml. These results agree with the water consumption data reported by Witter (1936). The water lost from the body, that eliminated from the cloaca plus that lost due to evaporation is presented in Table 4. As the water intake, the amount of water lost by all routes by a one-weekold chick was 20.0 gm. This water loss increased steadily as the birds grew, until at 32 weeks of age the amount the birds lost was on the average 406.9 gm. This figure also includes the water lost by the bird in the egg. The evaporative water loss has been reported by Medway and Kare (1957). Water balance has been worked out in the mature hen by Hart and Essex (1942) and Dixon (1958). In both cases the work revolved around the controversy of whether or not the cloaca plays a role in the reabsorption of water. Dixon (1958) exteriorized both the ureters and the rectum. In studying the water balance in birds, Dixon (1958) did not take into account the evaporative water loss which is quite appreciable; he only measured the water recovered in the urine and feces. Hart and Essex (1942) calculated the evaporative water loss. They obtained results ranging from 110-114 grams per day. They also state that since the amount of water lost through the skin is doubtless negligible these figures must represent water loss via the respiratory tract. The evaporative water loss that the author obtained for mature birds was approximately one half that reported by Hart and Essex (1942). The factor which probably plays the biggest role as far as altering the flow of urine is concerned is the handling of the bird. It

WATER METABOLISM AND BALANCE REFERENCES

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