- Email: [email protected]
Plasma antidiuretic hormone (ADH) concentrations in cattle, during various water and feed regimes
Cony
3i~l~/if~~. Plmid.
Vol. 81A.
No,
4. pp.
755-759,
0300-9629~85
1985
$3.00
+ 0.00
f‘, 1985 Pergamon Press Ltd
Printed in Great B&in
PLASMA ANTIDIURETIC HORMONE (ADH) CONCENTRATIONS IN CATTLE, DURING VARIOUS WATER AND FEED REGIMES* B. A.
BECKER,?
Department
of Dairy
M. B.
BOBER,$
Husbandry,
F. D.
University
(Recked
EL-NOUTY$
of Missouri,
and H. D. Columbia,
JOHNSON
MO 65201, USA
24 October 1984)
Abstract-l,
Twelve steers of three different breeds were exposed to five feed and water regimes in order to characterize changes in plasma antidiuretic hormone (ADH) concentrations. 2. No breed differences were found in plasma ADH concentration. 3. Plasma ADH concentration rose (4.2 to 22.0 pg/ml) during dehydration. 4. By 3 hr hydration, plasma ADH concentrations dropped dramatically (over 500~) to 9.2 pg/ml. 5. No changes in plasma ADH concentrations occurred during feed restriction and refeeding. 6. Hematocrit percentages were also determined and differences are hypothesized to relate to probable differences in environmental adaptability and genetic selection for meat or milk production among the three breeds. INTRODUCTION
et al., 1980). No such investigations have been reported in cattle. The objective of this study was to characterize plasma ADH in three breeds of cattle when feed and water were available ad libitum and during dehydration, hydration, feed restriction and refeeding. In order to find possible genetic differences, three diverse Bos taurus cattle breeds were selected. These were: (1) the Texas Longhorns, historically known for thriving under minimal management and nutritional conditions; (2) the Herefords, a European beef breed that performs well both in feedlots and under range conditions; and (3) the Holsteins, a dairy breed selected for milk production under good nutritional conditions and for good temperament.
Very little research has been done to characterize the function and role of the posterior pituitary antidiuretic hormone (ADH) in cattle. MacFarlane et ul. (1967) did comparative studies in sheep, cattle and camels, finding cattle the least sensitive to exogenous ADH. Johnson (1972) injected vasopressin in Bas taurus and Bos in&us crossbred cows and found no significant differences in renal response. ADH as measured by radioimmunoassay was found to decrease in Zebu cattle exposed to 39°C (Sief, 1973). This contrasted with later work by El-Nouty et al. (1980) who found an increase in ADH levels in Holstein cattle exposed to 35°C. ADH was found to increase in cattle when water was restricted to 50% for seven days (&if et al., 1973) and when dehydrated at 35-C for 30 hr (El-Nouty ef al., 1980). El-Nouty
et al. (1978) injected adrenocorticotropic hormone under both thermoneutrality (2OC) and heat (33C) and found ADH levels to increase within 5 min and remain significantly higher for 2 hr. All the above studies were concerned with the role of ADH in water balance and heat stress irrespective of diet. Nutritional interaction with ADH has been investigated in rats with results being very contradictory (Henrikx et ul., 1958; Radford, 19.59; Little and Radford,
1964; Forsling
et al., 1968; Kappagoda
*Contribution from Missouri Agr. Exp. Sta. Journal Series No. 9731. Approved by Director. Mention of trade names or companies does not constitute an implied warranty (or endorsement) by the University of Missouri or authors. TPresent address: Employed by the University of Nebraska at the Roman L. Hruska US. Meat Animal Research Center, ARS, USDA, P.O. Box 166, Clay Center, NE 68933, U.S.A. IDepartment of Physiological Sciences, School of Veterinary Medicine. Oklahoma State Ilniversity, Stillwater, OK 74078, U.S.A. aAnimal Production Unit, Faculty of Agriculture, Alcxandria University, Alexandria, Egypt,
MATERIALS AND METHODS
Experimental design and animals are described by Becker et al. (1985). To review brieflv. four Holsteins. four Herefords ‘and ‘four Texas Longhorn steers were randomly divided into two groups. Feed and water ud lib&urn were provided to all animals for 6 days and this is referred to as “the basal period”. Group one was then dehydrated for 54 hr followed by hydration at 48 hr. At the same time, group two was feed restricted to l/3 of basal period feed intake for 168 hr, followed by a 72-hr period with feed and water nd libitum (refeeding). After several days rest, all animals were subjected to a second basal period. Then the treatments were switched for each group such that group one was feed restricted and refed while group two was dehydrated and hydrated. Following a rest period, the experiment was completely replicated. Jugular catheters were inserted (Gomila and Roussel, 1976) in each steer several days before the experiment began. Blood sampling occurred as shown in Table 1. Twenty milliliters of blood was collected in EDTA-coated vacutainer tubes and immediately placed in ice. Samples were centrifuged at 86Og for 30 min and then plasma was frozen at - 2O’C until further analysis for antidiuretic hormone by radioimmunoassay (El-Nouty ef ai., 1978). Plasma hematocrit percents were determined immediately. Only samples from the first replicate were used for hormonal analysis. Basal samples were pooled for one compositie sample. Means were analyzed separately for each treatment (basal, dehydration, hydration, feed restriction and re755
8. A. BECKEK et u(.
756
Table I. Blood samplrng schedule Basal period Dehydration Hydration Feed restriction Refeedmg
0900, 1600 for last 3 days 0. I?. 24. 30, 36, 42. 48, 54 hr 3. 6, 12. 18.24, 36. 48, 90 hr 0. 24. 48. 72. 84, Y6. 108. 120. 132. 144. 156. 168 hr 6. 12. 18. 24, 36, 48%72 hr
feeding) by least-square analysis of variance as a split-plot by time with repeated measurements on the same animal (Gill and Hafs, 1971). Group and breed were on the main plot with group x breed as the error term. Time and all possible interactions were on the subplot. Orthogonal polynomial effects were tested for sjgni~cance. Least sjgnificance difference was used to test significance between means. Since hematocrit percentage during dehydration and hydration were significantly different between breeds, analysis of covariance was used to evaluate plasma ADH levels based on the assumption of all breeds having equal hematocrits. During all periods water intake. urine and fecal water content were monitored and have been reported elsewhere (Becker et al., 1985). To review, collections for water intake, urine output and fecal water content were made for 24-hr periods during the basal period, hydration, feed restriction and refeeding. During 54-hr dehydration collection periods were two 24-hr periods and one 6-hr period. The results for the mean of the basal periods and the final collection period for dehydration, hydration, feed restriction and refeeding are summarized in Table 2.
RESULTS
are shown in Fig. I. Plasma concentrations during the basal period were similar (P > 0.05) for all three breeds, 5.0 f 0.2, 4.2 f 0.3, 3.7 + 0.5 pg/ml for Texas Longhorn, Holstein and Hereford steers, respectively. Plasma ADH levels increased five-fold during 54-hr dehydration with the response being similar for all breeds. Mean concentrations rose from 4.2 pg/ml at 0 hour to 22.01 pg/ml at the end of 54 hr. During hydration no differences in breed responses were found. During this period, plasma ADH concentrations dropped extremely rapidly, over 50?,,, in the first 3 hr. By 12 hr plasma levels had declined to a mean 6.24pg/ml and did not significantly (P > 0.05)
change
thereafter.
Hematocrit During the basal period, hematocrit percentages were significantly higher in the Texas Longhorn than (P < 0.05) in the Holstein and Hereford steers (32.6 kO.2 vs 27.3 kO.7, and 29.8 +0.2x, respectively). Hematocrit percent (Table 3) differed (P ~0.05) Dehydration
Water intake, urine andj&cal water tjolume Changes in water intake, urine and fecal water volume have been reported elsewhere (Becker ef al., 1985) and are reviewed in Table 2. Briefly during the basal period, water intake was lower (P < 0.05) for the Texas Longhorn steers than the Holstein and Hereford steers. Urine and fecal water content were similar. During dehydration, hydration, feed restriction and refeeding, changes in all three parameters occurred with time but no breed differences were found. Basal period, ADH
dehydration,
and hydrution-plasma
Plasma antidiuretic hormone (ADH) concentrations during the basal period and changes in ADH levels during 54-hr dehydration and 4%hr hydration
Fig. 1. Plasma ADH (pgjml) concentrations 4 FIereford and 4 Holstein steers during rehydration.
in 4 Longhorn. dehydration and
Table 2. Summary of changes in water intake (liters). urine volume (liters), and fecal water content (liters) during the live dill&em feeding and water regimes for Texas Longhorn (TL), Hereford (H) and Holstein (HO) steers Water HO
TL 36.0 i
1.2”
49.3 + 0.88
H 46.1
+ 0.9” 0
TL 12.2 + 0.1 0.9 f 0.4
0
0
30.7 + 4.5
48.7 $: 4.5
40.1 k4.5
13.1 &-1.8
14.4 i: I.8
13.1 ir 1.8
8.2 & I.8
28.0 + 1.3
40.6 F 1.3
36.5 & I 3
9.6 & 0.8
II.1 kO.8
Urine HO Basal* 18.0 i: 2.5 Debydrationt 0.8 i: 0.4 Hydrationf 16.3 rt 0.8 Feed restrictions 6.2 + I.8 Refeeding/j 13.9 * 0.8
H
TL
Fecal water HO
H
20.6 2 3.0
15.5 f 0.3
17.5 * 0.2
16.6 i_ 0.4
0.8 + 0.7
0 8 i 0.7
0.8 * 0.7
17.5 f 0.8
12.1 kO.5
15.6 i 0.5
12.7 * 0.5
2.9 + 0 4
3.6 ir 0.4
4 6 i_ 0.4
12.3 i_ 0.6
IO.3 + 0.6
6.9 +
I .S
14.4 f 0.8
11.2+0.6
A%eans in the same row with common superscripts are not significantly different (P > 0.05). *Mean ( i SEM) for six days when feed and water were offered ad lihirum. Includes values for four hasal periods. tMean ( f SEM) for the last 6 hr of 54 hr of dehydration. :Mean ( f SEM) for the last 24 hr of 48 hr of hydration. $Medn ( f SEM) for the last 24 hr of 168 hr of feed restrictloo. lIMean ( _t SEM) for the last 24 hr of 72 hr of refeeding.
08107
ADH in dehydrated
among the three breeds during both dehydration and hydration. Overall hematocrit percentages were higher (P < 0.05) in the Texas Longhorn steers. The increase (P < 0.001) during dehydration was a moderate 3% (27-30.6%) in Holstein steers. In contrast, hematocrit percentages increased 13% in the Hereford steers (28-33x) and 17% in the Texas Longhorn steers (30-39”/,). At 3 hr hydration hematocrit percentages were actually higher than at any other time but then rapidly decreased. Analysis of covariance to evaluate the response of plasma ADH concentrations on the basis of equal hematocrits was significant (P < 0.01) only with respect to time. Adjusted means did not alter the magnitude of the responses. Feed restriction
and refeeding-plasma
ADH
Plasma ADH concentrations during feed restriction and refeeding are shown in Table 4. No changes (P > 0.05) due to treatment were found nor were there any differences (P > 0.05) in levels among the three breeds. Hematocrit
DISCUSSION
A major significance of this study is the documentation of an increase of plasma ADH concentrations for three breeds of cattle during 54-hr dehydration and the rapid decrease during hydration, which has not been previously reported. Becker et al. (1985) reported that the Longhorn steers tended to excrete less urine (Table 2) under the same conditions which led us to anticipate a higher plasma ADH concentration in the Texas Longhorn steers. The absence of
Table 3. Hematocrit percent ( f SEM) changes during 54-hr dehydration and 48.hr hydration of Texas Longhorn (TL), Hereford (H) and Holstein (HO) steers
0
Table 4. Plasma ADH concentrations (pg/ml + SEM) during feed restriction and refeeding in Texas Longhorn (TL), Hereford (H) and Holstein (HO) steers HOUS
0
24 48 72 84 96 108 120 132 I44 156 168 6 I2 I8 24 36 48 72
TL
HO
H
Feed restriction 4.4 * I .2 4.2 k 1.2 4.4 3.6 5.5 5.6 4.5 7.3 5.7 6.3 4.5 6.3 5.2 5.7 5.6 3.7 5.3 3.6 5.2 5.7 3.7 4.5 4.2 6.1 Refeeding 4.5 i 0.5 5.9 * 0.5 3.2 5.4 5.8 5.9 3.5 6.1 4.2 5.0 4.0 6.3 3.0 4.1
3.9 _t 1.2 2.5 2.9 3.2 1.9 2.6 2.4 3.8 3.4 3.9 2.4 4.5 3.3 * 0.5 2.8 2.7 2.1 3.5 4.2 3.4
percentages
Hematocrit percentages (Table 5) increased slightly (P < 0.05) during feed restriction. No significant (P > 0.05) changes occurred during hydration. No differences (P > 0.05) between breeds were found even though hematocrit percentages overall in the Texas Longhorn steers were greater (P > 0.05) during refeeding.
HOUS
757
and feed restricted steers
TL
12 24 30 36 42 48 54
30.0d.” + 0.8 32.6‘,” 33.W,” 36.9b.4 36.7b.4 37. Ih.” 38.6”.h.4 39.7”,A
6 12 18 24 36 48 90
37.1”,” 34.7” M 34.4”.M 33.50.M 33.3”M 31.0P.M 30.3p M
H Dehydration 28.2d.A + 0.8 2*,6c.d.L3 1,y.h.‘.R 3 I .8”,b,B 32.0d.b.B 29.6h.‘,” 33.4*,” 32.4*,’
30.9”,” 79 5”.“.N ;0:8”o 29.4w.p.N 29,3wN 28.7p.N 29.2p.“.Y
HO
difference between breeds in plasma ADH levels and the contrasting differences in urine output may be related to different rates of clearance for the breeds. ADH is metabolized via the kidney (66%) and the liver (33%) (Lauson et al., 1967), with 10% of the exogenous ADH appearing in the urine within 10 min and 28-100x appearing within 24 hr (Bauman and Dingham, 1976). If the changes in the glomerular filtration rate and plasma renal blood flow differed among these three breeds during dehydration as Johnson (1971) found in British and Zebu-crossed animals, then these plasma levels might not be truly indicative of relative biological activity. It is also possible that the Texas Longhorn steers which tended to have lower urine output (Table 2), are more sensitive to ADH. MacFarlane et al. (1967) found a difference in sensitivity to exogenous ADH among camels, sheep and cattle, but Johnson (1972) found no differences between Bos taurus and Bos indicus
Table 5. Hematocrit ::, during feed restriction and refeeding of Texas Longhorn (TL). Hereford (H), and Holstein (HO) steers HOUS
27.WA + 0.8 27,3kB28.36.h.C.B 29.6”.h.C 28,3”.b.‘.C 2*,7.&J & 30.6d,B 29.Y.’
30.5rI.0 28.9p.q.N 32.4m,h 29.5”.p-N 29.3~” 27.7q.N 28.7P.q ’
’ b.c.d.m~“.“P~qMezdns in the same column with common superscripts are not significantly different (P z 0.05). AB.C~M~“.oM~ansin the same row with common superscripts are not sigmficantly different (P > 0.05).
0 24 48 72 84 96 108 120 132 I44 I56 168 6 I2 I8 24 36 48 72
TL 31.2f0.7 32.0 31.3 34. I 32.7 32.0 32.7 33.1 33.5 32.6 33.0 33.6 35.3 + 0.8 35.6 33.6 35.2 35.3 33.6 35.9
H
HO
Feed restriction 28.2 & 0.7 27.7 i 0.7 29.9 30.4 29.9 29 .O 30.2 32.3 30.7 31.9 28.9 31.3 30.0 31.0 30.0 30.0 30.6 30.0 30.6 30.8 30.5 30.3 31.2 30.8 Refeeding 30.9 k 0.8 31.6iO.8 30.9 30.6 31.0 30.7 30.2 30. I 30.2 29.7 29.9 30.0 30.0 29.1
All breeds 29.0 f 0.4 30.8 30.0 32.2 31.7 30.7 31.2 31.0 31.4 31.3 31.3 31.8 32.6 i 0.8 32.2 31.8 31.8 31.7 31.2 31.7
758 cattle. There also exists the third possibility
8. A. BECKEK e1 al
that there is no difference in any of these factors (absolute plasma levels, clearance rate or sensitivity) and that the difference in urine output without a corresponding breed difference in plasma ADH may be due to anatomical differences in the kidney. Schoen (1969) found that animals that were capable of concentrating their urine had a greater percentage of the kidney occupied by the medulla. Plasma ADH concentration did not change during feed restriction or refeeding. Sensitivity to exogeneous ADH has been found to vary with types of diet (Kappagoda er ai., 1980) leading us to investigate possible changes with level of feed intake. However, Becker et al. (1985) showed that during feed restriction plasma osmolality significantly fluctuated between 268 and 260mOs/kg and that during refeeding plasma osmolality peaked at 278 mOs/kg at 6 hr and then dropped to 268 mOs/kg by 24-hr refeeding. These changes in plasma osmolahty should have been physiologically significant enough to stimulate changes in plasma ADH concentrations. The sensitivity of the radioimmunoassay for ADH was 0.08 pg/tube (El-Nouty et a/., 1978) which would be able to detect the theoretical decrease of 3.4 pg/ml ADH for the one unit decrease in plasma osmolality. In humans. for each increment in plasma osmoIality, an increase of plasma ADH on the average of 0.34 pg/ml has been found (Robertson, 1974; Robertson et a/., 1976). However, also in humans a threshold above which a small increase will elevate ADH secretion and below which there is no inhibition is reported to be 280 mOs (Robertson. 1974; Robertson et ul., 1976). If there is a similar threshold in cattle, then the plasma levels under these conditions can be assumed to be below the threshold level. Changes in hematocrit percentages during dehydration and hydration were significantly different among the three breeds. The magnitude of increase in the Hereford (13”/0) and Longhorn (170;,) is similar to that found in other beef steers (Bianca ei al., 1965; Weeth rt al., 1967; Rumsey and Bond, 1976) and the overall lower hematocrits and smaller increase (3?;) in the Holstein steers are similar to results reported for Ayrshires dehydrated for three days (Bianca, 1970). The higher hematocrit values at 36-hr hydration is thought to be the result of excitement associated with offering of water after a period of dehydration, because animals rehydrated by infusion through a rumen cannula have been reported not to show this response (Bianca. 1970). Interpretations of these breed differences in hematocrit percentages are difficult. The differences may be related to differences in adaptability, as seen in the hematological comparisons between Brahman and Hereford cattle in a warm climate (Turner, 1980). Perhaps the Holstein steers are able to maintain plasma volume, indicated by the small increase in hematocrit percent in the same way as the camel (Schmidt-Nielsen, 1964). In contrast, maybe the failure to maintain plasma volume, as seen so clearly in the Texas Longhorn steers, is an adaptive mechanisms of tolerance, similar to the camel’s ability to tolerate wide fluctuations in body temperature (Schmidt-Nielsen ef (I/.. 1957). Or possibly there is a difference related to the type of production in dairy
versus beef breeds, i.e. the Holsteins have lower hematocrits, which may in turn reflect a greater plasma volume, thereby insuring necessary water for milk production. However, no significant breed differences were found during feed restriction and refeeding even though hematocrit percentages in the Texas Longhorn steers tended to be higher, particularly during refeeding. This may mean that under a certain range of conditions breed responses are similar; however, differences may be found when the breeds are confronted with certain stresses (like dehydration). This would support the idea that the differences are associated with adaptability versus genetic differences related to production, In conclusion, plasma ADH concentration was not found to differ among Texas Longhorn, Herefords and Holstein steers. In the bovine plasma ADH increased five-fold during 54-hr dehydration and decreased rapidly (50% within the first 3 hr) upon hydration. Plasma ADH concentrations were not affected by level of feed intake.
REFERENCES
Bauman G. and Dingham J. (1976) Distribution,
blood transport, and degradation of antidiuretic hormone in man. j. c&t. Invest. 57, 1109-l 116. Becker B.. Bober M., El-Nouty F. and Johnson H. (1985) Prolactin and its involvement in fluid regulation in the bovine. Comp. Biochem. Physiol. 81A, 93-98. Bianca W. (1970) EfTects of dehydration, rehydration, and overhydration on the blood and urine in oxen. Br. IX’,. .1. 126, 121.-131. Bianca W., Finlay J. and McLean J. (1965) Response of steers to water restriction. Res. CEI. Sri, 6, 38-54. El-Nouty F., Elbanna I., Davis T. and Johnson H. (1980) Aldosterone and ADH response to heat and dehydration in cattle. J. appl. Physiol. 48, 249-255. El-Nouty F., Elbanna, I. and Johnson H. (1978) E&t ot adrenocorticotropic hormone on plasma glucocorticords and antidiuretic hormone of cattle exposed to 20 and 33°C. 3. Dairy Sci. 61, 189-196. Forsling M., Jones J. and Lee J. (1968) Factors influencing the sensitivity of the rat to vasopressin. J. P/~~sio/. 196, 4955505. Gill J. and Hafs H. (lY71) Analysis of repeated measurements of animals. J. Anim. SC;. 33, 3311336. Gomila L. and Roussel J. (I 976) A modification of iueular cannulation technique. j. Dair.F Sci. 59, 24. i Hendrikx A. and Epstein F. (1958) Effect of feeding protcm and urea on renal concentrating ability in the rat. ,4117.J. Phvsiol. 195, 539-542. Johnson K. (1971) Renal function in Bos /atlru.s and Bos in&us-crossbred COWSunder conditions of normal hydration and mild hydration. Rrv. t’e~. Sci. 12, 438-447. Johnson K. (1972) The effect of vasopressin on urinary excretion of Bos ~rrurus and &IS in[~i~ff.s-~rossbr~iCOWS. Res. (‘et. Sci. 13, 431-435. Kappagoda C., Linden R., and Pashiey M. (1980) Increased sensitivity of ADH bioassay in rats by change in diet. J. Physiol. 299, 4251135. Lauson H. (1967) Metabolism of antidiuretic hormone. .&t. J. Med. 42, 113-744. Little J. and Radford E. Jr. (1964) Bioassay for antidiurcli~ activity in blood of undistur~d rats. J. uppl. Pbwioi. 19, 1799186. MacFarlane W., Kinne R.. Walmsley C., Siebert B. and Peter D. (I 967) Vasopressin and the increase of water and electrolyte excretion by sheep. cattle, and camels. Nn(urc 214, 979-98 I.
ADH
in dehydrated
and feed restricted
Radford E. (1959) Factors modifying water metabolism in rats fed dry diets. Am. J. Physiol. 196, 1098-1108. Robertson G. (1974) Vasopressin in osmotic regulation in man. A. Rea. Med. 23, 315-321. Robertson G.. Shelton R. and Athar S. (1976) The osmoregulation of vasopressin. Kidney Int. i0, 25-27. Rumsey T. and Bond J. (1976) Cardiorespiratory patterns, rectal temperature, serum electrolytes, and packed cell volume in beef cattle deprived of feed and water. .I. Anim. Sci. 42, 1227-1237. Schmidt-Nielson K. (1964) Desert Animals, pp. 63-67. Oxford University Press, London, Schmidt-Nielsen K:, Schmidt-Nielsen B., Jarnon S. and Hoypt T. (1957) Body temperature of the camel and its relation to water economy. Am. J. Physiol. 188, 103-l 12.
steers
759
Schoen A. (1969) Water conservation and the structure of the kidneys of tropical bovids. J. Physiol. 204, 143P-144P. Seif S. (1973) Water kinetics, heat adaptability and ADH of bovine. Ph.D. Thesis, University of Missouri. Seif S., Johnson H. and Hahn L. (1973) Environmental heat and partial water restriction effects on body fluid spaces, water loss, body temperature, and metabolism of Holstein cows. J. Diary Sci. 56, 581-586. Turner J. (1980) Genetic and biological aspects of Zebu adaptability. J. Anim. Sci. 50, 1201-1205.. Weeth H., Sawtiney D. and Lesperance A. (1967) Changes in body fluids, excreta, and kidney function of cattle deprived water. J. Anim. Sri. 26, 418423.
3i~l~/if~~. Plmid.
Vol. 81A.
No,
4. pp.
755-759,
0300-9629~85
1985
$3.00
+ 0.00
f‘, 1985 Pergamon Press Ltd
Printed in Great B&in
PLASMA ANTIDIURETIC HORMONE (ADH) CONCENTRATIONS IN CATTLE, DURING VARIOUS WATER AND FEED REGIMES* B. A.
BECKER,?
Department
of Dairy
M. B.
BOBER,$
Husbandry,
F. D.
University
(Recked
EL-NOUTY$
of Missouri,
and H. D. Columbia,
JOHNSON
MO 65201, USA
24 October 1984)
Abstract-l,
Twelve steers of three different breeds were exposed to five feed and water regimes in order to characterize changes in plasma antidiuretic hormone (ADH) concentrations. 2. No breed differences were found in plasma ADH concentration. 3. Plasma ADH concentration rose (4.2 to 22.0 pg/ml) during dehydration. 4. By 3 hr hydration, plasma ADH concentrations dropped dramatically (over 500~) to 9.2 pg/ml. 5. No changes in plasma ADH concentrations occurred during feed restriction and refeeding. 6. Hematocrit percentages were also determined and differences are hypothesized to relate to probable differences in environmental adaptability and genetic selection for meat or milk production among the three breeds. INTRODUCTION
et al., 1980). No such investigations have been reported in cattle. The objective of this study was to characterize plasma ADH in three breeds of cattle when feed and water were available ad libitum and during dehydration, hydration, feed restriction and refeeding. In order to find possible genetic differences, three diverse Bos taurus cattle breeds were selected. These were: (1) the Texas Longhorns, historically known for thriving under minimal management and nutritional conditions; (2) the Herefords, a European beef breed that performs well both in feedlots and under range conditions; and (3) the Holsteins, a dairy breed selected for milk production under good nutritional conditions and for good temperament.
Very little research has been done to characterize the function and role of the posterior pituitary antidiuretic hormone (ADH) in cattle. MacFarlane et ul. (1967) did comparative studies in sheep, cattle and camels, finding cattle the least sensitive to exogenous ADH. Johnson (1972) injected vasopressin in Bas taurus and Bos in&us crossbred cows and found no significant differences in renal response. ADH as measured by radioimmunoassay was found to decrease in Zebu cattle exposed to 39°C (Sief, 1973). This contrasted with later work by El-Nouty et al. (1980) who found an increase in ADH levels in Holstein cattle exposed to 35°C. ADH was found to increase in cattle when water was restricted to 50% for seven days (&if et al., 1973) and when dehydrated at 35-C for 30 hr (El-Nouty ef al., 1980). El-Nouty
et al. (1978) injected adrenocorticotropic hormone under both thermoneutrality (2OC) and heat (33C) and found ADH levels to increase within 5 min and remain significantly higher for 2 hr. All the above studies were concerned with the role of ADH in water balance and heat stress irrespective of diet. Nutritional interaction with ADH has been investigated in rats with results being very contradictory (Henrikx et ul., 1958; Radford, 19.59; Little and Radford,
1964; Forsling
et al., 1968; Kappagoda
*Contribution from Missouri Agr. Exp. Sta. Journal Series No. 9731. Approved by Director. Mention of trade names or companies does not constitute an implied warranty (or endorsement) by the University of Missouri or authors. TPresent address: Employed by the University of Nebraska at the Roman L. Hruska US. Meat Animal Research Center, ARS, USDA, P.O. Box 166, Clay Center, NE 68933, U.S.A. IDepartment of Physiological Sciences, School of Veterinary Medicine. Oklahoma State Ilniversity, Stillwater, OK 74078, U.S.A. aAnimal Production Unit, Faculty of Agriculture, Alcxandria University, Alexandria, Egypt,
MATERIALS AND METHODS
Experimental design and animals are described by Becker et al. (1985). To review brieflv. four Holsteins. four Herefords ‘and ‘four Texas Longhorn steers were randomly divided into two groups. Feed and water ud lib&urn were provided to all animals for 6 days and this is referred to as “the basal period”. Group one was then dehydrated for 54 hr followed by hydration at 48 hr. At the same time, group two was feed restricted to l/3 of basal period feed intake for 168 hr, followed by a 72-hr period with feed and water nd libitum (refeeding). After several days rest, all animals were subjected to a second basal period. Then the treatments were switched for each group such that group one was feed restricted and refed while group two was dehydrated and hydrated. Following a rest period, the experiment was completely replicated. Jugular catheters were inserted (Gomila and Roussel, 1976) in each steer several days before the experiment began. Blood sampling occurred as shown in Table 1. Twenty milliliters of blood was collected in EDTA-coated vacutainer tubes and immediately placed in ice. Samples were centrifuged at 86Og for 30 min and then plasma was frozen at - 2O’C until further analysis for antidiuretic hormone by radioimmunoassay (El-Nouty ef ai., 1978). Plasma hematocrit percents were determined immediately. Only samples from the first replicate were used for hormonal analysis. Basal samples were pooled for one compositie sample. Means were analyzed separately for each treatment (basal, dehydration, hydration, feed restriction and re755
8. A. BECKEK et u(.
756
Table I. Blood samplrng schedule Basal period Dehydration Hydration Feed restriction Refeedmg
0900, 1600 for last 3 days 0. I?. 24. 30, 36, 42. 48, 54 hr 3. 6, 12. 18.24, 36. 48, 90 hr 0. 24. 48. 72. 84, Y6. 108. 120. 132. 144. 156. 168 hr 6. 12. 18. 24, 36, 48%72 hr
feeding) by least-square analysis of variance as a split-plot by time with repeated measurements on the same animal (Gill and Hafs, 1971). Group and breed were on the main plot with group x breed as the error term. Time and all possible interactions were on the subplot. Orthogonal polynomial effects were tested for sjgni~cance. Least sjgnificance difference was used to test significance between means. Since hematocrit percentage during dehydration and hydration were significantly different between breeds, analysis of covariance was used to evaluate plasma ADH levels based on the assumption of all breeds having equal hematocrits. During all periods water intake. urine and fecal water content were monitored and have been reported elsewhere (Becker et al., 1985). To review, collections for water intake, urine output and fecal water content were made for 24-hr periods during the basal period, hydration, feed restriction and refeeding. During 54-hr dehydration collection periods were two 24-hr periods and one 6-hr period. The results for the mean of the basal periods and the final collection period for dehydration, hydration, feed restriction and refeeding are summarized in Table 2.
RESULTS
are shown in Fig. I. Plasma concentrations during the basal period were similar (P > 0.05) for all three breeds, 5.0 f 0.2, 4.2 f 0.3, 3.7 + 0.5 pg/ml for Texas Longhorn, Holstein and Hereford steers, respectively. Plasma ADH levels increased five-fold during 54-hr dehydration with the response being similar for all breeds. Mean concentrations rose from 4.2 pg/ml at 0 hour to 22.01 pg/ml at the end of 54 hr. During hydration no differences in breed responses were found. During this period, plasma ADH concentrations dropped extremely rapidly, over 50?,,, in the first 3 hr. By 12 hr plasma levels had declined to a mean 6.24pg/ml and did not significantly (P > 0.05)
change
thereafter.
Hematocrit During the basal period, hematocrit percentages were significantly higher in the Texas Longhorn than (P < 0.05) in the Holstein and Hereford steers (32.6 kO.2 vs 27.3 kO.7, and 29.8 +0.2x, respectively). Hematocrit percent (Table 3) differed (P ~0.05) Dehydration
Water intake, urine andj&cal water tjolume Changes in water intake, urine and fecal water volume have been reported elsewhere (Becker ef al., 1985) and are reviewed in Table 2. Briefly during the basal period, water intake was lower (P < 0.05) for the Texas Longhorn steers than the Holstein and Hereford steers. Urine and fecal water content were similar. During dehydration, hydration, feed restriction and refeeding, changes in all three parameters occurred with time but no breed differences were found. Basal period, ADH
dehydration,
and hydrution-plasma
Plasma antidiuretic hormone (ADH) concentrations during the basal period and changes in ADH levels during 54-hr dehydration and 4%hr hydration
Fig. 1. Plasma ADH (pgjml) concentrations 4 FIereford and 4 Holstein steers during rehydration.
in 4 Longhorn. dehydration and
Table 2. Summary of changes in water intake (liters). urine volume (liters), and fecal water content (liters) during the live dill&em feeding and water regimes for Texas Longhorn (TL), Hereford (H) and Holstein (HO) steers Water HO
TL 36.0 i
1.2”
49.3 + 0.88
H 46.1
+ 0.9” 0
TL 12.2 + 0.1 0.9 f 0.4
0
0
30.7 + 4.5
48.7 $: 4.5
40.1 k4.5
13.1 &-1.8
14.4 i: I.8
13.1 ir 1.8
8.2 & I.8
28.0 + 1.3
40.6 F 1.3
36.5 & I 3
9.6 & 0.8
II.1 kO.8
Urine HO Basal* 18.0 i: 2.5 Debydrationt 0.8 i: 0.4 Hydrationf 16.3 rt 0.8 Feed restrictions 6.2 + I.8 Refeeding/j 13.9 * 0.8
H
TL
Fecal water HO
H
20.6 2 3.0
15.5 f 0.3
17.5 * 0.2
16.6 i_ 0.4
0.8 + 0.7
0 8 i 0.7
0.8 * 0.7
17.5 f 0.8
12.1 kO.5
15.6 i 0.5
12.7 * 0.5
2.9 + 0 4
3.6 ir 0.4
4 6 i_ 0.4
12.3 i_ 0.6
IO.3 + 0.6
6.9 +
I .S
14.4 f 0.8
11.2+0.6
A%eans in the same row with common superscripts are not significantly different (P > 0.05). *Mean ( i SEM) for six days when feed and water were offered ad lihirum. Includes values for four hasal periods. tMean ( f SEM) for the last 6 hr of 54 hr of dehydration. :Mean ( f SEM) for the last 24 hr of 48 hr of hydration. $Medn ( f SEM) for the last 24 hr of 168 hr of feed restrictloo. lIMean ( _t SEM) for the last 24 hr of 72 hr of refeeding.
08107
ADH in dehydrated
among the three breeds during both dehydration and hydration. Overall hematocrit percentages were higher (P < 0.05) in the Texas Longhorn steers. The increase (P < 0.001) during dehydration was a moderate 3% (27-30.6%) in Holstein steers. In contrast, hematocrit percentages increased 13% in the Hereford steers (28-33x) and 17% in the Texas Longhorn steers (30-39”/,). At 3 hr hydration hematocrit percentages were actually higher than at any other time but then rapidly decreased. Analysis of covariance to evaluate the response of plasma ADH concentrations on the basis of equal hematocrits was significant (P < 0.01) only with respect to time. Adjusted means did not alter the magnitude of the responses. Feed restriction
and refeeding-plasma
ADH
Plasma ADH concentrations during feed restriction and refeeding are shown in Table 4. No changes (P > 0.05) due to treatment were found nor were there any differences (P > 0.05) in levels among the three breeds. Hematocrit
DISCUSSION
A major significance of this study is the documentation of an increase of plasma ADH concentrations for three breeds of cattle during 54-hr dehydration and the rapid decrease during hydration, which has not been previously reported. Becker et al. (1985) reported that the Longhorn steers tended to excrete less urine (Table 2) under the same conditions which led us to anticipate a higher plasma ADH concentration in the Texas Longhorn steers. The absence of
Table 3. Hematocrit percent ( f SEM) changes during 54-hr dehydration and 48.hr hydration of Texas Longhorn (TL), Hereford (H) and Holstein (HO) steers
0
Table 4. Plasma ADH concentrations (pg/ml + SEM) during feed restriction and refeeding in Texas Longhorn (TL), Hereford (H) and Holstein (HO) steers HOUS
0
24 48 72 84 96 108 120 132 I44 156 168 6 I2 I8 24 36 48 72
TL
HO
H
Feed restriction 4.4 * I .2 4.2 k 1.2 4.4 3.6 5.5 5.6 4.5 7.3 5.7 6.3 4.5 6.3 5.2 5.7 5.6 3.7 5.3 3.6 5.2 5.7 3.7 4.5 4.2 6.1 Refeeding 4.5 i 0.5 5.9 * 0.5 3.2 5.4 5.8 5.9 3.5 6.1 4.2 5.0 4.0 6.3 3.0 4.1
3.9 _t 1.2 2.5 2.9 3.2 1.9 2.6 2.4 3.8 3.4 3.9 2.4 4.5 3.3 * 0.5 2.8 2.7 2.1 3.5 4.2 3.4
percentages
Hematocrit percentages (Table 5) increased slightly (P < 0.05) during feed restriction. No significant (P > 0.05) changes occurred during hydration. No differences (P > 0.05) between breeds were found even though hematocrit percentages overall in the Texas Longhorn steers were greater (P > 0.05) during refeeding.
HOUS
757
and feed restricted steers
TL
12 24 30 36 42 48 54
30.0d.” + 0.8 32.6‘,” 33.W,” 36.9b.4 36.7b.4 37. Ih.” 38.6”.h.4 39.7”,A
6 12 18 24 36 48 90
37.1”,” 34.7” M 34.4”.M 33.50.M 33.3”M 31.0P.M 30.3p M
H Dehydration 28.2d.A + 0.8 2*,6c.d.L3 1,y.h.‘.R 3 I .8”,b,B 32.0d.b.B 29.6h.‘,” 33.4*,” 32.4*,’
30.9”,” 79 5”.“.N ;0:8”o 29.4w.p.N 29,3wN 28.7p.N 29.2p.“.Y
HO
difference between breeds in plasma ADH levels and the contrasting differences in urine output may be related to different rates of clearance for the breeds. ADH is metabolized via the kidney (66%) and the liver (33%) (Lauson et al., 1967), with 10% of the exogenous ADH appearing in the urine within 10 min and 28-100x appearing within 24 hr (Bauman and Dingham, 1976). If the changes in the glomerular filtration rate and plasma renal blood flow differed among these three breeds during dehydration as Johnson (1971) found in British and Zebu-crossed animals, then these plasma levels might not be truly indicative of relative biological activity. It is also possible that the Texas Longhorn steers which tended to have lower urine output (Table 2), are more sensitive to ADH. MacFarlane et al. (1967) found a difference in sensitivity to exogenous ADH among camels, sheep and cattle, but Johnson (1972) found no differences between Bos taurus and Bos indicus
Table 5. Hematocrit ::, during feed restriction and refeeding of Texas Longhorn (TL). Hereford (H), and Holstein (HO) steers HOUS
27.WA + 0.8 27,3kB28.36.h.C.B 29.6”.h.C 28,3”.b.‘.C 2*,7.&J & 30.6d,B 29.Y.’
30.5rI.0 28.9p.q.N 32.4m,h 29.5”.p-N 29.3~” 27.7q.N 28.7P.q ’
’ b.c.d.m~“.“P~qMezdns in the same column with common superscripts are not significantly different (P z 0.05). AB.C~M~“.oM~ansin the same row with common superscripts are not sigmficantly different (P > 0.05).
0 24 48 72 84 96 108 120 132 I44 I56 168 6 I2 I8 24 36 48 72
TL 31.2f0.7 32.0 31.3 34. I 32.7 32.0 32.7 33.1 33.5 32.6 33.0 33.6 35.3 + 0.8 35.6 33.6 35.2 35.3 33.6 35.9
H
HO
Feed restriction 28.2 & 0.7 27.7 i 0.7 29.9 30.4 29.9 29 .O 30.2 32.3 30.7 31.9 28.9 31.3 30.0 31.0 30.0 30.0 30.6 30.0 30.6 30.8 30.5 30.3 31.2 30.8 Refeeding 30.9 k 0.8 31.6iO.8 30.9 30.6 31.0 30.7 30.2 30. I 30.2 29.7 29.9 30.0 30.0 29.1
All breeds 29.0 f 0.4 30.8 30.0 32.2 31.7 30.7 31.2 31.0 31.4 31.3 31.3 31.8 32.6 i 0.8 32.2 31.8 31.8 31.7 31.2 31.7
758 cattle. There also exists the third possibility
8. A. BECKEK e1 al
that there is no difference in any of these factors (absolute plasma levels, clearance rate or sensitivity) and that the difference in urine output without a corresponding breed difference in plasma ADH may be due to anatomical differences in the kidney. Schoen (1969) found that animals that were capable of concentrating their urine had a greater percentage of the kidney occupied by the medulla. Plasma ADH concentration did not change during feed restriction or refeeding. Sensitivity to exogeneous ADH has been found to vary with types of diet (Kappagoda er ai., 1980) leading us to investigate possible changes with level of feed intake. However, Becker et al. (1985) showed that during feed restriction plasma osmolality significantly fluctuated between 268 and 260mOs/kg and that during refeeding plasma osmolality peaked at 278 mOs/kg at 6 hr and then dropped to 268 mOs/kg by 24-hr refeeding. These changes in plasma osmolahty should have been physiologically significant enough to stimulate changes in plasma ADH concentrations. The sensitivity of the radioimmunoassay for ADH was 0.08 pg/tube (El-Nouty et a/., 1978) which would be able to detect the theoretical decrease of 3.4 pg/ml ADH for the one unit decrease in plasma osmolality. In humans. for each increment in plasma osmoIality, an increase of plasma ADH on the average of 0.34 pg/ml has been found (Robertson, 1974; Robertson et a/., 1976). However, also in humans a threshold above which a small increase will elevate ADH secretion and below which there is no inhibition is reported to be 280 mOs (Robertson. 1974; Robertson et ul., 1976). If there is a similar threshold in cattle, then the plasma levels under these conditions can be assumed to be below the threshold level. Changes in hematocrit percentages during dehydration and hydration were significantly different among the three breeds. The magnitude of increase in the Hereford (13”/0) and Longhorn (170;,) is similar to that found in other beef steers (Bianca ei al., 1965; Weeth rt al., 1967; Rumsey and Bond, 1976) and the overall lower hematocrits and smaller increase (3?;) in the Holstein steers are similar to results reported for Ayrshires dehydrated for three days (Bianca, 1970). The higher hematocrit values at 36-hr hydration is thought to be the result of excitement associated with offering of water after a period of dehydration, because animals rehydrated by infusion through a rumen cannula have been reported not to show this response (Bianca. 1970). Interpretations of these breed differences in hematocrit percentages are difficult. The differences may be related to differences in adaptability, as seen in the hematological comparisons between Brahman and Hereford cattle in a warm climate (Turner, 1980). Perhaps the Holstein steers are able to maintain plasma volume, indicated by the small increase in hematocrit percent in the same way as the camel (Schmidt-Nielsen, 1964). In contrast, maybe the failure to maintain plasma volume, as seen so clearly in the Texas Longhorn steers, is an adaptive mechanisms of tolerance, similar to the camel’s ability to tolerate wide fluctuations in body temperature (Schmidt-Nielsen ef (I/.. 1957). Or possibly there is a difference related to the type of production in dairy
versus beef breeds, i.e. the Holsteins have lower hematocrits, which may in turn reflect a greater plasma volume, thereby insuring necessary water for milk production. However, no significant breed differences were found during feed restriction and refeeding even though hematocrit percentages in the Texas Longhorn steers tended to be higher, particularly during refeeding. This may mean that under a certain range of conditions breed responses are similar; however, differences may be found when the breeds are confronted with certain stresses (like dehydration). This would support the idea that the differences are associated with adaptability versus genetic differences related to production, In conclusion, plasma ADH concentration was not found to differ among Texas Longhorn, Herefords and Holstein steers. In the bovine plasma ADH increased five-fold during 54-hr dehydration and decreased rapidly (50% within the first 3 hr) upon hydration. Plasma ADH concentrations were not affected by level of feed intake.
REFERENCES
Bauman G. and Dingham J. (1976) Distribution,
blood transport, and degradation of antidiuretic hormone in man. j. c&t. Invest. 57, 1109-l 116. Becker B.. Bober M., El-Nouty F. and Johnson H. (1985) Prolactin and its involvement in fluid regulation in the bovine. Comp. Biochem. Physiol. 81A, 93-98. Bianca W. (1970) EfTects of dehydration, rehydration, and overhydration on the blood and urine in oxen. Br. IX’,. .1. 126, 121.-131. Bianca W., Finlay J. and McLean J. (1965) Response of steers to water restriction. Res. CEI. Sri, 6, 38-54. El-Nouty F., Elbanna I., Davis T. and Johnson H. (1980) Aldosterone and ADH response to heat and dehydration in cattle. J. appl. Physiol. 48, 249-255. El-Nouty F., Elbanna, I. and Johnson H. (1978) E&t ot adrenocorticotropic hormone on plasma glucocorticords and antidiuretic hormone of cattle exposed to 20 and 33°C. 3. Dairy Sci. 61, 189-196. Forsling M., Jones J. and Lee J. (1968) Factors influencing the sensitivity of the rat to vasopressin. J. P/~~sio/. 196, 4955505. Gill J. and Hafs H. (lY71) Analysis of repeated measurements of animals. J. Anim. SC;. 33, 3311336. Gomila L. and Roussel J. (I 976) A modification of iueular cannulation technique. j. Dair.F Sci. 59, 24. i Hendrikx A. and Epstein F. (1958) Effect of feeding protcm and urea on renal concentrating ability in the rat. ,4117.J. Phvsiol. 195, 539-542. Johnson K. (1971) Renal function in Bos /atlru.s and Bos in&us-crossbred COWSunder conditions of normal hydration and mild hydration. Rrv. t’e~. Sci. 12, 438-447. Johnson K. (1972) The effect of vasopressin on urinary excretion of Bos ~rrurus and &IS in[~i~ff.s-~rossbr~iCOWS. Res. (‘et. Sci. 13, 431-435. Kappagoda C., Linden R., and Pashiey M. (1980) Increased sensitivity of ADH bioassay in rats by change in diet. J. Physiol. 299, 4251135. Lauson H. (1967) Metabolism of antidiuretic hormone. .&t. J. Med. 42, 113-744. Little J. and Radford E. Jr. (1964) Bioassay for antidiurcli~ activity in blood of undistur~d rats. J. uppl. Pbwioi. 19, 1799186. MacFarlane W., Kinne R.. Walmsley C., Siebert B. and Peter D. (I 967) Vasopressin and the increase of water and electrolyte excretion by sheep. cattle, and camels. Nn(urc 214, 979-98 I.
ADH
in dehydrated
and feed restricted
Radford E. (1959) Factors modifying water metabolism in rats fed dry diets. Am. J. Physiol. 196, 1098-1108. Robertson G. (1974) Vasopressin in osmotic regulation in man. A. Rea. Med. 23, 315-321. Robertson G.. Shelton R. and Athar S. (1976) The osmoregulation of vasopressin. Kidney Int. i0, 25-27. Rumsey T. and Bond J. (1976) Cardiorespiratory patterns, rectal temperature, serum electrolytes, and packed cell volume in beef cattle deprived of feed and water. .I. Anim. Sci. 42, 1227-1237. Schmidt-Nielson K. (1964) Desert Animals, pp. 63-67. Oxford University Press, London, Schmidt-Nielsen K:, Schmidt-Nielsen B., Jarnon S. and Hoypt T. (1957) Body temperature of the camel and its relation to water economy. Am. J. Physiol. 188, 103-l 12.
steers
759
Schoen A. (1969) Water conservation and the structure of the kidneys of tropical bovids. J. Physiol. 204, 143P-144P. Seif S. (1973) Water kinetics, heat adaptability and ADH of bovine. Ph.D. Thesis, University of Missouri. Seif S., Johnson H. and Hahn L. (1973) Environmental heat and partial water restriction effects on body fluid spaces, water loss, body temperature, and metabolism of Holstein cows. J. Diary Sci. 56, 581-586. Turner J. (1980) Genetic and biological aspects of Zebu adaptability. J. Anim. Sci. 50, 1201-1205.. Weeth H., Sawtiney D. and Lesperance A. (1967) Changes in body fluids, excreta, and kidney function of cattle deprived water. J. Anim. Sri. 26, 418423.