Brit. uet. ]. (1964),
120,
6g
WATER DIURESIS IN CATTLE By
RoDGER
G.
DALTON
Department of Veterinary Medicine, University of Glasgow
The diuretic response which follows the oral administration of water has been extensively studied in monogastric animals but little is known about the diuretic response after ingestion of water in ruminant animals. Klisiecki, Pickford, Rothschild & Verney (I 932) have shown that, in man and dog, the diuretic response following ingestion of water is mainly influenced by the time required for the absorption of water from the intestine and the effects of the antidiuretic hormone. They also showed that ingested water passed rapidly through the stomach into the intestine and that the diuretic response was not affected by retention of water in the stomach. Little is known about diuresis in the cow or other ruminant animals. Andersson ( I955) gave goats water loads equivalent to 20 per cent of their body weights and did not observe any adverse effects. He found that the diuretic response in these animals lasted over I 3 hours. From this study he concluded that since a large volume of water could be administered orally to goats without producing water intoxication, water absorption must be slower in goats than in non-ruminant animals. The present work was undertaken to determine the diuretic response following the oral administration of water to cows. MATERIALS AND METHODS
Experiments were carried out on 24 healthy Ayrshire cows whose body weights ranged from 45o-6oo kilograms. Twelve of the cows were lactating, the others were not. The cows were housed and were fed twice daily. Four of the cows were fed a total daily ration of approximately 8o lb. cut grass, 7-12 lb. hay and 4-12 lb. concentrated cattle cake. The other 20 cows were fed a total daily ration of appro,ximately 40 lb. turnips, 30 lb. silage, 7- I2 lb. hay and 4-12 lb. concentrated cattle cake. The amounts fed were adjusted according to the requirements of each animal. The cows had access to water before and during the experiments, although in fact none of the cows was observed to drink during an experiment. Normal variations in urine flow during the day were measured in four cows and the diuretic response to water loading was measured in one of these four animals and in the 20 other cows. These experiments were started in the morning, about an hour after the cows had been fed and milked, and were continued until shortly before the cows were fed and Inilked in the evening. In the diuresis experiments, urine was collected for a period of one to two hours before the administration of the water load. The experiments were conducted
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with minimal disturbance to the cows and their behaviour during the experiments was little different from that of other cows in the byre. Urine was collected using a self-retaining urethral catheter. The catheter consisted of a soft rubber tube 50 em. long and I em. diameter, with an i.t;l.flatable cuff halfway along its length. In one half of the catheter were multiple circular perforations go mm. in diameter. This multi-perforate part of the catheter and the inflatable cuff were inserted into the bladder and the cuff was filled with 50 to I oo ml. of water. The catheter was then retracted so that the inflatable cuff lodged in the neck of the bladder. A long drainage tube was attached to the exposed end of the catheter and the bladder was drained by continuous siphonage. To facilitate insertion of the · catheter and to alleviate irritation and consequent straining when it was in situ, light posterior epidural anaesthesia was maintained by the periodic injection of2 to 4 ml. of 5 per cent procaine hydrochloride solution. A water load of I8 litres was administered in all the diuresis experiments. The water was administered with a pump through a thin rubber stomach tube passed via the nostril into the oesophagus until the tip was estimated to be at the cardia. The water was heated to approximately 39 oc before administration. The time taken to insert the stomach tube and administer the water ranged from 3-7 minutes. Urine osmolalities were determined with a Hortvet Cryoscope on urine samples collected from two cows. RESULTS
Normal variations in urine flow Table I shows, for each of four cows, the mean and standard deviations of TABLE I NORMAL VARIATIONS IN URINE FLOW DURING THE DAY
~
~A
~B
22 "3 12"3 I2"3 I7 "3 I4"0 11"0 8·7 8·7 9·6 9 ·6 9 ·6 15"3 8 ·o 17"3 19"3 8·6 I6· 5 6·o
I7"7 19"3 13"3 I0·6 I2"0 I3"0 I0 ·8 8·7 9"0 9"0 8·8 7"3 8 ·8 8·o 7"5 4"0 I2"5 7"9 7"3 I2"9
I2 ·6 ±5"I
I0•4 ±3"5
~C
~D
12"7 II"9 6·6 9"0 8·7 14"4 7"3 11 "0 II ·O 11"0 JI•O 7"7 11"5 13"3 10"0 11 "3 8•7 9"3 7"3
12"9 11"1 I0·8 8·o 8 ·8 7"7 8·g 7"2 10"0 4 "0 8·8 s·o 8·3
ml.fmin.
II.OO a.m. I I.I5 I 1.30 II.45 I2.00 noon I2"I5 I2.30 I2.45 1.00 p.m. I.I5 1.30 1·45 2.oop.m. 2.I5 2"30 2"45 3.00 p.m. 3-I5 3"30 3-45
Mean S.D.
I0·2 ±2·2
8·6 ±2"5
WATER DIURESIS IN CATTLE
urine flow per minute when determined at IS-minute intervals during the day. The rate of urine flow of each of these cows was not constant during the day, as is shown by the size of the standard deviations relative to the mean values. From these results it was assumed that an increase in urine flow of more than twice the mean value was outside the range of normal variations. In the diuresis experiments an increase in urine flow of more than twice that before water loading is therefore described as a diuretic response.
Diuretic response to water loading In Table II are shown the mean urine flows per minute of 2I cows when determined at IS-minute intervals over a period of one to two hours before administration of water loads. All the cows showed diuretic responses following water loading, but variations occurred in the time which elapsed between water loading and the onset of diuresis and in the time taken to reach peak diuresis. Table II shows for each cow the time which elapsed before the onset of diuresis, the time taken to reach peak diuresis, and the maximal urine flow observed. TABLE II THE DIURETIC RESPONSE IN CATTLE
Cow D E F G H I
J
K L M N 0
p
Q
R
s u
T
v
w X
Mean S.D.
Urineflow (ml.fmin.) Maximum Bifore after loading loading 12 "5 14"3 17"5 g·2 21•1 11·g 10"5 5"9 8·1 5"1 g ·6 11"5 10•3 14"7 13"5 6·8 13 "1 g·l 5"g 11 •3 15"7 11"3 ±4"0
Minutes after water loading To To Start of Maximum diuresis* diuresis
13g 123 lOg 103 g8 100 105 g3 113 llg 63 164 107 14g 117 23 g6 85 127 78 143
6o 6o 45 15 30 6o 6o 15 45 15 120 15 75 15 45 30 6o 30 30 15 30
75 150 150 75 6o 255 210 135 120 150 240 225 105 go 165 225 105 210 go 135 75
107"3 ±3o·g
45t ±30
150t ±6o
*Diuresis was assumed to occur when the urine flow was twice the mean preloading urine flow. t Calculated to the nearest 15-minute interval.
The time before the onset of diuresis after water loading ranged from IS to I 20 minutes, with a mean value of 4S ± go minutes. In I 3 of the 2 I cows diuresis occurred within a period of 30 to 6o minutes after water loading; in Q
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six cows it occurred within I 5 minutes of water loading, and in two others more than 75 minutes after water loading. Peak diuresis occurred on the average 150 ± 6o minutes after water loading, with a range of 6o to 255 minutes. In seven of the 2 I cows peak diuresis occurred within I20 to I8o minutes after loading; in eight cows it occurred within I20 minutes of loading, and in six cows not until more than 180 minutes after water loading. The mean maximal urine flow was approximately ten times greater than the mean urine flow before water loading. Examples of the types of diuretic responses observed in cows are shown in Figs. 1, 2, 3, 4 arid 5· From these figures it can be seen that there was a marked variation in the types of diuretic response. Figs. I, 2 and 4 show examples of the response of some cows in which the urine flow increased rapidly to a peak after water loading and then gradually declined. Approximately half the cows showed a phasic diuretic response similar to those shown in Figs. 3 and 5· It is of interest to note how the changes in urine flow shown in Figs; 4 and 5 were accompanied by changes in urine osmolality. 20
200
~IS
i Figs.
1
and
2.
0 50
6.
Diuretic response to water loading in a cow.
0-D
~-\/'-\"0&.$/LITC I \.~l,-,
ISO
20
0·1 O· Z.
0
<
r o.
O·J
9
Oo+
I 0
O·S
o-6
I.
&.
I.
+.
HOUP.S.
Figs. 3 and 4· Diuretic response to :water loading in a cow.
~.
"-
WATER DIURESIS IN CATTLE
ISO
0·1
0
c
~
73
I
r;.. ~~r;·
Fig. 5· Diuretic response to water loading in a cow.
One animal, cow S, had a very poor diuretic response. In this animal the urine flow did not rise above 23 ml.Jmin. and very little of the administered water load was excreted within six hours. Since the normal variations in urine flow during the day were small in comparison to the urine flow during diuresis, the extent of the diuretic response was calculated from the difference in the urine output before and after water loading. For each cow the difference between the successive half-hourly urine output after water loading and the mean half-hourly urine output before water loading was calculated. In Table III these differences are expressed as a TABLE III PERCENTAGE WATER LOAD EXCRETED AT HALF-HOUR INTERVALS
Minutes after water Loading Cow D E F G H I
J
K L
M N 0
p
Q
R
s
T
u v w X
Mean S.D.
30 1'4 o·5 o·5 3·6 4"4 o·o o·o 2 "1 0•7 1"7 0"9 3"5 0"7 5"2 1"0 1"3 0"4 J·8 3"0 3"2 3"7
6o 8·8 2•2 4 "8 11"1 12"0 2 "1 2 "0 4"4 3"0 2"5 o·o 3"1 o·5 13"2 3 "9 1·1 2"9 2•2 11·6 7"2 19"2
go 20 '3 10•0 12"5 12•1 g·8 7"9 6 ·5 g·6 8·7 12"3 o·o o·5 6·9 21 "4 8·9 0 "1 11•2 6·6 18· 5 7•8 15 ·o
. 120 17 "0 16 ·3 14•1 7"3 4 "8 5"3 10 "4 13 ·8 16 ·5 17 •8 1"0 2 "1 15"8 12"3 15·1 0"7 11"3 8· 1 15"2
JO·B
5"9
150 8·1 17"3 14'8 6·2 8·7 6·g 12 "7 11·8 12"0 18·1 3"0 13·8 10"3 10"9 16·7 0"7 7"4 9"9 12"5 10"2 10"4
ISO
210
7"9 15·8 11·6 5"3 4"8 3 "9 12 "1 9"2 8 ·5 15"0 4"7 18·5 6·1 7 "9 15·6 o ·5 5"9 9 "4 3 "3 8 ·3 2 "9
4"7 7"9 7"5 8 ·4 6 ·o 6 ·7 15·o 9 "3 7"0 11"9 5"2 5"2 4"6 5·o 13·8 o ·6 4"0 12"3 6·1 8·6 5·8
240 2"5 2"2
270 3"4
300 3"9
330 3·6
13·8 6 ·1 7•8 13·8 6·2 8·2 5"7 8·6 16·3 2•2 4 "2 10"9 1"9 o·9 7"7 9"7 6·4 3"3
6·7 3"6 12 ·8 11 "3 9 "4 11 "5 6·1 4"5 4"7 1"7 3"9 7"2 1"0 0 "4 4 "2 9 "2 1·5 4"2
o ·1 0"2 5"1 7"9 10"4 1o·g 3·6 5"5 2"7 0"4 0"3 2•2 1"4 o·7 1"4 2"0 o·1 1·5
0"4 o ·o 3"2 6 ·5 7"2 6 ·5 4 "4 5"9 1"7 0"4 o ·9 0"7 0"4 5 ·8 3 "8 0"4 1·0
g6o
o ·o 1"0 3 "2 6·5 2 "5 6·2
0 "2 0 "3 o ·o 3"9 2 ·6 o·o 0 '5
2 "1 6 ·9 g·S J0•6 I0·6 8·4 s-6 3"2 2 "9 7"4 5 ·6 1"9 ±1·5 ±5 · 1 ±5'9 ±s-6 ±4'4 ±4·8 ±3 '5 ±4'4 ±3 ' 7 ±3'4 ±2 ·6 ±2 '3
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120, 2
percentage of the water load. Table IV shows the cumulative percentage water loads excreted. Fig. 6 shows the mean percentage water and Fig. 7 the cumulative mean percentage water load excreted by the cows at successive half-hourly intervals after water loading. From the results shown in Tables III and IV and in Figs. 6 and 7, it can be seen that there was a wide range in the percentage water load excreted. The average diuretic response was not complete six hours after water loading, by which time 7 r per cent of the water load had been excreted. TABLE IV CUMULATIVE PERCENTAGE WATER LOAD EXCRETED AT HALF-HOUR INTERVALS
Minutes after water loading Cow D E F
G H I
1
K L M N
0
p
Q R
s T
u
v w X
Mean S.D.
30 I •4 o·s o·s 3·6 4•4 o·o o·o 2•1
0•7 1•7 o·g 3 ·s 0•7 5"2 1•0 1•3 0•4 1·8 3•0 3•2 3•7
6o 1o·2 2·7 s·3 14•7 !6•4 2·1 2 ·o 6·5 3·7 4·2 o·g 6·6 1•2 18·4 4·9 2·4 3•3 4·o 14·6 10•4 22·9
go 3o ·s 12·7 17 ·8 26·8 26·2
w·o 8·s 16·1 12·4 16·s o·g 7·1 8·1 39·8 13·8 2·5 14•5 10·6 33·1 18·2 37•9
120 47 ·5 29·0 31·9 34•1 31 •0 '5"3 18·9 29·9 28·9 34·3 1·9 g·2 23•9 52·1 28·g 3·2 25 ·8 18·7 48·3 2g·o 43·8
150 ss-6 46·3 46·7 40•3 39•7 22·2 31·6 41·7 4o·g 52·4 4·9 23·o 34•2 63·o 4s-6 3·9 33•2 28·6 6o·8 39 •2 54·2
180 63 ·5 62·1 s8·3 45"6 44·s 26·1 43·7 50·9 49·4 67·4 g·6 41·s 40•3 7o·g 61·2 4·4 39•1 38·o 64·1 47•5 57•1
210 68·2 70·0 6s-8 54•0 so·s 32·8 s8·7 60·2 s6·4 79·3 14·8 46·7 44•9 75·9 7s·o yo 43•1 50·3 70·2 56 •1 62·9
240 70·7 72·2
270 74·1
300 78·o
330 81·6
67·8 s6·6 40·6 72·5 66·4 64·6 8s·o 23·4 63·o 47•1 8o·1 8s·g 6·g 44•0 58·o 79·9 62·5 66•2
74•5 6o·2 s3·4 83·8 75"8 76·1 91·1 27·9 67·7 48·8 84·o 93·1 7·9 44.4 62 2 8g·1 64·0 70·4
74•6 6o·4 s8·s 91"7 86·2 87·o 94·7 33·4 70·4 49•2 84·3 95·3 g·3 45·1 63·6 91·1 64·1 71 •9
75"0 6o·4 6o·4 61·7 62·7 g8·2 93•4 g6•6 93·5 1oo·o gg·1 101·6 39·3 45·5 72·1 49·6 g6·2 JO·o 45"5 6g·4 94·9 64·5 72 •9
36o
g6·4 10·3 45.5 73·3 97·5 64·5 73·4
I ·g 7·5 17·3 27·9 38·5 46·g 54·3 6o·7 6s ·7 68·g 71·o 71·4 ±•·s ±6· 4 ±11•1 ±13•7 ±1 5 ·g ±16·7 ±19·2 ±20·1 ±22·0 ±22·8 ±19•2 ±27•4 dO.
...
0
WATER DIURESIS IN CA"LE.
..."'
!;(
~15.
*-' /
j.
..."'
.ar/1
i
I
z
o•o.
..."'::> ...... 5 5.
,
:t,
·-........... . . . . .:,.. ..... +
• ....-.
/ T' :: •I *.-··.- - - .,. i-
/
z
"'
/
··-·-<. .
4
····-...
•
~ ·-....
,-'
... -..
-4-
·--•• ,
-4~
····-:!(l
~
· :,-._ ..,~
.,.__
• ......... '• ,..._ __ ........
,
0 L-~~~---T----~----~----~--~ L 2. 3. 4. 5. 6.
H2!UU
Fig. 6. Percentage water load excreted (see also Table III).
WATER DIURESIS IN CATTLE
75
100
~90 1-
~ao u X ... 70 0
~ 60 ...J
Fig. 7· Cumulative percentage water load excreted (see also Table IV).
Effect of lactation on diuretic response To determine whether the amount of the water load excreted was affected by lactation, the cows were divided into three groups depending on their milk production. Table V shows the percentage water load excreted two, four and TABLE V RELATIONSHIP OF PERCENTAGE WATER LOAD EXCRETED AND MILK YIELD
Percentage water load excreted after loading Cow D G H
J
K
M
Q
R
w
lb. milk/day 0 0
0 0 0 0 0 0
0
Mean S.D. F I N
0
p
20 6 20 10 13
Mean S.D. E L
s
T
u v X
Mean S.D.
45 40 34 31 35 33 33
2 hours 47'2 34'I 3I'0 I9·o 30'0 34'3 52'I 27'0 29'0
4 hours 70 '4 67 ·8 56·6 72'5 66 ·5 85 ·o 8o·o 83·9 62 ·5
6 hours
6o ·5 g6 •7 100 94'5 64'5
33'7 ±9·8
71•8 ±9'7
83'2 ±19·1
32'0 I4'9 1'9 9'1 23'9
40'3 23'4 62·9 47 ' 1
62'3 45'6
16·4 ±11·9
43'4 ±16·4
53'9 ± 11·8
28·9 29'0 3'2 25'8 18·7 48'2 43·8
72'1 64 •7 6 ·8 44'1 57 '9 79·8 66·2
100 10'4 45'6 73'1 97'2 73'5
28·2 ±15'1
55 '9 ±24'4
66·6 ± 34'0
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six hours after water loading by nine cows which were not lactating, five cows producing up to 2 gallons per day, and seven cows producing over 3 gallons per day. The mean percentage water load excreted two, four and six hours after water loading by the non-lactating cows was greater than that excreted by the lactating cows (apart from two hours after water loading, when the five cows producing up to 2 gallons per day had excreted more than the non-lactating cows). On statistical analysis it was found, however, that there was no significant difference in the mean percentage water load excreted two, four and six hours after water loading by the non-lactating and the lactating cows .
.
DISCUSSION The results obtained in this study show that there is a wide variation in the type of diuretic response following oral administration of water to cattle. There is no statistical correlation between these variations and lactation. It is of interest to compare the diuretic response observed in cattle with that described in the dog. Wolf (Igso) described how diuresis in the dog after water loading is characterized by a delay of I 8 to 42 minutes before the onset of diuresis, followed by a rapid rise in urine flow to peak values and a subsequent short period of continued diuresis, leading to the elimination of an amount of water approximately equal to the water load administered. Smith (Igso) states that in the dog diuresis does not begin until 40 minutes after water loading and the maximum diuresis is not reached until so minutes or more after water loading. He states that the average time after water loading to peak diuresis in five dogs was 102 minutes. Neither of these authors describes a phasic variation in the diuretic response in dogs comparable to that observed in cattle in this study. In this study it was found that in cattle the average time after water loading to the onset of diuresis was 4S ± 30 minutes. (range I5-I20 minutes) and to peak diuresis was ISO ± 6o minutes (range 6o-2ss minutes). Adolph (1943) calculated that in the dog the time tin hours required to complete the diuretic response to water is related to the water load LH 2o (in percentage of body weight) by the equation t = I "3 + LH20 In this study the water load (r8litres) was equivalent to 3 to 4 per cent ofthe body weights of the cows (4so-6oo kg.). Using the equation formulated by Adolph the time taken for a dog to excrete a water load equivalent to 3 to 4 per cent body weight is 4 ·3 to s ·3 hours. In this study the average diuretic response in cattle lasted more than six hours. From this study it was concluded that in cattle there was considerable individual variation in the type of diuretic response to oral water loading. The diuretic response in cattle was also more protracted than that described in the dog. The differences observed in the diuretic response following oral administration of water to cattle and in that described in the dog may be associated with differences in the rates of water absorption from the alimentary tract in the two species.
WATER DIURESIS IN CATTLE
77
REFERENCES
ADoLPH, E. F. (I943)· Physiological Regulations. Lancaster, Pa.: Cattell. ANDERSSON, B. (I955). Acta physiol. scand., 33, I. KLISIECKI, A., PICKFORD, M., RoTHSCHILD, P. & VERNEY, E. B. (I932). Proc. roy. Soc. B., 112,496. SMITH, H. W. (rgso). The Kidney. New York: Oxford University Press. WoLF, A. V. (I950). The Urinary Function of the Kidney. New York: Grune & Stratton. ·
(Received for publication I 9 July, I 963)