Journal of Physiology (1989) 417, pp.

421-435 421
With 7 text-ftgure8
Printed in Great Britain

EFFECTS OF DEHYDRATION AND REHYDRATION ON

THERMOREGULATORY SWEATING IN GOATS
BY M. A. BAKER
From the Division of Biomedical Sciences, University of California,
Riverside, CA 92521, USA
(Received 28 February 1989)
SUMMARY
1. Measurement of rectal temperature (Tr) sweat rate, respiratory frequency (I)
and respiratory evaporation (Eresp) were made in one Nubian and four Alpine-
Toggenberg goats while they stood for 90 min in a climate chamber at 40 0C ambient
temperature (T,). The animals were studied when they were hydrated, when they had
been dehydrated by 48 h water deprivation, and when they were rehydrated by
voluntary drinking of water or saline or by intraruminal water administration.
Plasma osmolality (Posm), plasma protein concentration (PP) and haematocrit (Hct)
were measured before every experiment and before and after voluntary drinking.
2. Hydrated animals increased evaporation by panting and sweating during heat
exposure and Tr rose about 1 0C. The rate of sweating was as high or higher than
Eresp. Dehydrated animals had lower sweat rates and higher Tr than hydrated
animals, butf and Eresp were the same in hydrated and dehydrated animals.
3. When dehydrated goats were allowed to drink after 60 min of heat exposure,
sweating began abruptly within 3 min of the start of drinking in every animal
whether water or saline was drunk. Sweat rate returned to hydrated levels or higher
before any change occurred in P.sm, PP or Hct. Respiratory frequency was higher
after drinking than in dehydrated animals which were not allowed to drink.
4. When water was administered by rumen tube after 60 min of heat exposure,
sweating in the Nubian occurred with a short latency, similar to the onset after
drinking. In the other four animals, sweating onset occurred on average at 13 min
42 s after intraruminal water administration.
5. It is concluded that sweating is a significant avenue of evaporative heat loss in
these goats when they are hydrated and exposed to high T.. Sweat rate is markedly
reduced after water deprivation but returns to hydrated levels within 3 min after the
start of drinking. The rapid recovery of sweating after voluntary drinking is not
initiated by changes in Posm or in blood volume and does not appear to depend upon
osmoreceptors in the mouth or gastrointestinal tract since it occurs after drinking
either water or saline. The arrival of water in the rumen may be sufficient to initiate
immediate sweating in some goats, but the act of drinking is necessary in others.
422 M. A. BAKER

INTRODUCTION

Dehydrated mammals in hot environments can save water by reducing the rate of
panting or sweating and regulating body temperature above hydrated levels
(Schmidt-Nielsen, Schmidt-Nielsen, Jarnum & Houpt, 1957; Taylor, 1970; Baker,
1984). Two consequences of dehydration, an elevation in the concentration of
solutes in body fluids and a reduction in the blood volume, can both influence
thermoregulatory water losses.
Evidence that elevated body fluid solute concentration is an important stimulus
for the thermoregulatory adjustments occurring in dehydration comes from studies
in which hypertonic solutions are infused into hydrated animals. When body fluid
osmolality is elevated by intragastric or intravenous administration of hypertonic
solution in heat-stressed mammals, thermoregulatory evaporation is reduced and
body temperature rises (Nielsen, 1974; Baker & Doris, 1982). A number of
observations support the possibility that intracranial osmoreceptors play a role in
this reduction of evaporative heat loss. Thus. thermal panting in hydrated dogs and
rabbits (Baker & Dawson, 1985; Turlejska & Baker, 1986) and sweating in monkeys
(Owen, Matthes & Gisolfi, 1987) are inhibited by intracarotid or intracerebro-
ventricular infusions of hypertonic solutions. The effects of isotonic reduction
of blood volume on thermoregulatory water losses have been studied in only a few
experiments. Reduced blood volume leads to reduced thermoregulatory water losses
and elevated body temperatures in exercising humans and resting rats (Fortney,
Nadel, Wenger & Bove, 1981; Horowitz & Meiri, 1985).
Observations in dehydrated humans and dogs have shown that thermoregulatory
evaporation may recover rapidly after drinking; however, the signals which initiate
this are not understood. Senay & Christiansen (1965) and Nicolaidis (1970) found
that dehydrated humans in a warm environment began to sweat within seconds to
minutes after drinking, but the relationships between plasma osmolality, blood
volume and sweating onset were not measured. In dehydrated dogs, we have recently
observed a rapid recovery of thermal panting after drinking which is not dependent
on changes in plasma osmolality or plasma volume (Baker & Turlejska, 1989).
The purpose of the present study was to investigate the thermoregulatory effects
of dehydration and rehydration in the goat, an animal which uses both panting and
sweating to control the body temperature in hot environments. The experiments
were designed to answer the following questions: how does water deprivation affect
body fluid osmolality, blood volume and thermoregulatory evaporation in goats?
What is the pattern of recovery of evaporative heat loss after drinking and how is it
related to osmolality of drinking fluid, osmolality of body fluids and blood volume'?
Does the act of drinking provide signals which affect thermoregulatory evaporation?
In these experiments, measurements of body temperature and of evaporation by
panting and sweating have been made in goats exposed to heat when they were
hydrated and dehydrated and when they were rehydrated by voluntary drinking of
water or saline or by the introduction of water into the rumen through a tube.
 RECO0VER Y OF SVVEATINTG AFTER DRINKING LINV GOATS 423

METHODS
Five adult castrated male goats were studied. Four of the animals were cross-bred
Alpine--Toggenbergs and one was a Nubian. They were purchased from a local goat farm one year
before these experiments began. At the farm they had been castrated four days after birth and had
been kept in outdoor corrals. After their arrival at this research facility, the goats were trained to
ride to the laboratory in a cart, to stanid quietly in a climate chamber and to wear a respiratory
mask. They were housed in covered outdoor (corrals during the day and were brought indoors at
night. They were fed alfalfa hay and mixed grain at 08-30 and 16 30 h and had free access to water
except during periods of dehydration. At the time that these studies began, the Alpine-Toggenberg
goats were 2 years old and weighed 50+4 kg (rnean+S.E.M.) and the Nubian was 3 years old and
weighed 7.15) kg. General procedures
Experiments started at 13.00 h. Before each experiment, the animal was brought to the
laboratory in a cart, prepared for blood sampling and for measurement of sweat rate, rectal
temperature (Tr) and respiratory frequency (f) and was then led into a walk-in temperature-
controlled chamber where it stood with its neck in a loose wooden stock. A window in the chamber
allowed the goat and the experimenter to see each other. Measurements were begun 15 min after
the animal entered the chamber (time 0 for data analysis). Air temperature (Ta) inside the chamber
was controlled at 40+ 1 'C. Relative humidity was measured but was not controlled and varied
between 23 and 70 % but did not change more than 10 % during a single experiment.
Each animal was studied in five separate experiments: (1) when it was hydrated ad libitum; (2)
when it was dehydrated after 48 h of water deprivation; (3) during water drinking after
dehydration; (4) during saline drinking after dehydration; and (5) during water administration by
rumen tube after dehydration.
Experimental protocols
In experiments without drinking, a single blood sample was taken by external jugular
venipuncture before the animal went into the chamber. In the drinking experiments, a 20 gauge x
2 in Teflon catheter (Delmed A-Cath) was inserted into the external jugular vein under local
anaesthesia (lidocaine hydrochloride) and was attached to a length of extension tubing which was
closed with a stopcock and taped to the back of the neck to allow blood sampling before and after
drinking.
Measurements were made for 90 min in experiments on hydrated and dehydrated animals
(experiments 1 and 2). In drinking and rumen tube experiments (experiments 3, 4 and 5),
measurements on the dehydrated animals were made for 60 min and then they drank or received
water by rumen tube, and measurements were continued for 30 min after the end of drinking.
Sweat rate and Tr were measured continuously during heat exposure in all experiments.
Respiratory frequency was measured every 5 min for a period of 1 min and respiratory evaporation
(Eresp) was measured every 15 min for a period of 5 min in the 90 min hydrated and dehydrated
experiments.
In the drinking studies, an experimenter entered the chamber at 60 min, drew a blood sample
from the indwelling catheter, and then offered the goat a bucket of warm (36 °C) tap water or of
NaCl solution. Preliminary experiments had indicated that the effect on sweating was the same if
the animal drank cool (21 °C) or warm fluid, and 36 'C was selected so that the fluid itself would
not cause a change in body temperature. Osmolality of the NaCl drinking fluid was adjusted to the
plasma osmolality (Posm) of each animal measured before the experiment. The goat was allowed to
drink to satiation and then the bucket was removed and the time spent drinking and volume drunk
were recorded. In the rumen tube studies, three experimenters entered the chamber at 60 min. One
held the goat's head, a second inserted a flexible plastic tube (9 mm o.d.) into the rumen through
the mouth and then the third pumped water (36 °C), equal to the weight loss of the animal during
the period of dehydration. from a bucket into the rumen.
To find out whether the sight of water would initiate sweating, at the end of the 90 min
dehydrated without drinking experiment a bucket of water was placed in front of two of the goats
without allowing them to drink.
424 M. A. BAKER
Measurement techniques
The rate of sweating was measured using the ventilated capsule technique described by Bullard
(1962). A plastic capsule (7 cm diameter) taped securely to the side of the neck was ventilated at
2-7 1 min-' with compressed air. Relative humidity and temperature of air entering and leaving the
capsule were measured with Vaisala humidity and temperature probes and recorded on a
rectilinear flat-bed recorder (Soltek). Average sweat rates were calculated after computer averaging
of the change in water content of air flowing through the capsule.
Chamber T. and Tr of the goat were measured using copper-constantan thermocouples and a
Bailey model BAT-12 thermocouple thermometer. Respiratory frequency was measured with a
strain gauge fastened to a tube stretched around the animal's chest and was recorded on a Grass
Model 7 polygraph. Respiratory evaporation was measured using an open-circuit system and a
plastic mask lined with rubber foam which was placed over the nose and mouth of the animal and
strapped behind the ears. Air was pulled through the mask with a calibrated vacuum motor and
water content of air entering and leaving the mask was monitored using resistance hygrometers
(American Instrument Company) as described previously (Baker, 1984). For measurements of
Eresp, an experimenter entered the chamber, placed the mask on the animal, and stayed in the
chamber for the 5 min measurement period.
A 3 ml blood sample was drawn into a heparinized syringe before each experiment and, in the
drinking experiments, additional samples were taken immediately before and at 5, 15 and 30 min
after the end of drinking. Haematocrit (Hct) was measured in triplicate using capillary tubes and
a microhaematocrit centrifuge. After centrifugation of the remainder of the sample, plasma was
separated from red cells, Posm was measured with a Wescor Model 5100 B vapour pressure
osmometer and total plasma protein concentration (PP) with an American Optical hand-held
refractometer. Changes in plasma volume will affect both Hct and PP; but PP probably provides
a more accurate index of plasma volume shifts because of the possibility that splenic red cell storage
in the goat varies with the behaviour of the animal as it does in sheep (Turner & Hodgetts, 1959).
Statistical analyses
All measurements are expressed as the mean± standard error of the mean (S.E.M.). Data were
analysed using two-way analysis of variance, analysis of variance for replicate measures (Sokol &
Rohlf, 1981) and paired t tests. The null hypothesis was rejected at the 0'05 % level.

RESULTS

Thermoregulation in hydrated and dehydrated animals

Hydrated animali
During the 90 min period of heat exposure, hydrated goats increased evaporation
by both panting and sweating and Tr rose about 1 °C (Fig. 1). In the first part of the
heat exposure, the rate of cutaneous moisture loss was low and similar to values
reported for passive diffusion of water across the skin (Dmi'el, Robertshaw &
Choshniak, 1979). The onset of sweating was marked by a sudden increase in
cutaneous moisture loss which occurred at 24X6 + 4X0 min in hydrated animals. The
pattern of sweating in these animals consisted of peaks of cutaneous moisture loss as
has been reported previously for goats (Robertshaw, 1968) and, once sweating had
begun, an increase in the level of moisture loss occurring between the peaks (Fig. 2,
top). Sweat rate changed significantly during the 90 min heat exposure (P < 0.001 by
two-way ANOVA). The Nubian sweated at rates which were 2-3 times the rate of
sweating of the other four animals, and this accounts for the large S.E.M. in sweat
rates in the figures where data from the five animals is pooled. For example, from
60-90 min, in the experiments shown in Fig. 1, the mean rate of sweating of the four
cross-bred goats was 74+11 g m-2 h-1 while the Nubian sweated at 170 g m-2h-1.
 RECOVERY OF SWEATING AFTER DRINKING IN GOATS 425

K uE
.0
4-

-0

CL
V IN

Lua CD,E

CD

a)

o 15 30 45 60 75 90
Time (min)
Fig. 1. Effect of dehydration on rectal temperature (Tr), respiratory frequency (f),
respiratory evaporation (Eresp) and sweat rate in five goats at Ta 40 'C. Each animal was
studied first when it was hydrated (@ for Tr,f and Eresp, open bars for sweat rate) and again
when it was dehydrated (O and hatched bars).

Dehydrated animals
During the period of water deprivation preceding the experiments on dehydrated
animals shown in Fig. 1, the goats lost 3-5 + 0-6 kg of body weight and Posm, Het and
PP all increased significantly (Table 1).
When the animals were dehydrated, f and Eresp were the same as in the hydrated
animals (insignificant difference by replicate measures ANOVA) but Tr was higher
(P < 0-001) and sweat rate was reduced (P < 0-001; Fig. 1). In three of the
426 M. A. BAKER
dehydrated animals, there were no sweat peaks during 90 min in the hot chamber
(Fig. 2, bottom, shows a record of cutaneous moisture loss from one of these animals).
In each of the other two animals, single peaks of sweating occurred at 79 and at
85 min, respectively. After these peaks, moisture loss returned quickly to low levels.

150-
100
50

Tco.
M -E

cnm

100 L
501_
i I I I I

15 30 45 60 75 90
Time (min)
Fig. 2. Cutaneous moisture loss (sweat rate) in an Alpine-Toggenberg goat at Ta 40 °C
when it was hydrated (top) and when it was dehydrated (bottom). Each trace shows the
output of a resistance hygrometer in the effluent airstream from a ventilated skin capsule.
The abscissa shows the time after beginning measurements. There were no sweat peaks
before 15 min.

Effects of drinking on thermoregulation

Water drinking
The period of water deprivation preceding the water drinking experiments resulted
in a loss of body weight of 3-6 + 0-4 kg and elevations in Posm and PP (Table 1).
Average rates of cutaneous moisture loss were low and steady before drinking (Fig. 3
and Fig. 4, top) and there was no change in sweat rate during the first 60 mmn of
heat exposure (two-way ANOVA). In three animals, there were no sweat peaks
before drinking. A single peak occurred in each of the other two animals, one at
40 min and one at 49 min. When the dehydrated animals were offered water, they
drank 3.9 + 0-3 1. The period of drinking was 80 + 10 s.
Every animal showed a peak of sweating after drinking water, and the latency to
the first sweat peak was 152 + 12 s after the beginning of drinking (Fig. 4, top). Sweat
rate changed significantly (P < 0-001) during the first 5 min period following
drinking (Fig. 3). Every animal had a higher sweat rate during the 30 min after
drinking water than during the last 30 min of the hydrated heat exposure, but this
was not statistically significant. The highest sustained sweat rate observed in any of
these experiments exceeded 300 g m-2 h-1 in the Nubian during the 30 min after
drinking water (Fig. 4, top). Body temperature tended to stabilize after drinking and
the average change in Tr from 60 to 90 min was significantly different (P < 0.05 by
paired t test) from the change in dehydrated animals who did not drink. At the end
 RECOVERY OF SWEATING AFTER DRINKING IN GOATS 427
of drinking, f was below the pre-drinking rate. It then rose above pre-drinking f and
average f during the 30 min after drinking was higher (P < 0-05) than f during the
last 30 min of heat exposure in dehydrated animals that did not drink. The highest
f observed in any animal in these studies (320 per minute) occurred 10 min after
drinking water. There was no change in Posm, Hct or PP from the time immediately
preceding drinking to 30 min after drinking (Table 1).

TABLE 1. Effects of dehydration and rehydration on plasma osmolality (P..m), haematocrit (Hct)
and plasma protein concentration (PP) in five goats
After drinking
Before Before
Hydrated heating drinkingt 5 min 15 min 30 min
Poem (mosmol(kg H20-')) 289+2
Hct(%) 34+1
PP(g'-1) 79+1
Dehydrated
PoSm 302+ 1**
Hct 39+ 1*
PP 87 +2**
Dehydrated
(H20 drinking)
Posm 300 +2*** 300+2 301+2 301+2 300+3
Hct 36+2 33+1 34+1 32+1 34+1
PP 86+2* 85+2 87+2 85+2 85+2
Dehydrated
(NaCi drinking)
PoSm 300+ 1* 299+1 298+ 1 300+1 300+1
Hct 36+1 34+1 36+2 34+1 33+1
PP 86+ 1** 84+1 85+1 84+1 82+1±
Dehydrated
(H20 by stomach tube)
p 0m 301+2*
Hct 38+1**
PP 86+2**
t At 60 min.
* Significantly different from hydrated values (paired t test).
t Significantly different from pre-drinking value (paired t test).
* or t = P < 005; ** = P < 0 01; *** = P < 01001.

Saline drinking
The period of water deprivation preceding the saline drinking experiments led to
a loss of body weight of 45 + 0-4 kg and elevations in Posm and PP (Table 1).
Cutaneous moisture loss did not change significantly during the 60 min before
drinking saline (Fig. 4, bottom and Fig. 5). Only one of the five animals showed a
sweat peak before drinking, and this occurred at 25 min, followed by a return of
cutaneous moisture loss to a low level. When offered NaCl solution to drink, the
dehydrated animals drank 5-1 + 0 4 1. The period of drinking was 147 + 6 s.
Every animal showed a peak of sweating after drinking saline (Fig. 4, bottom) and
the latency to the first sweat peak was 149±14 s after the beginning of drinking.
428 M. A. BAKER
Sweat rate changed significantly (P < 0-0001) in the first 5 min period following
drinking (Fig. 5). The average change in Tr during the 30 min period after drinking
NaCl was not significantly different from the change during the same period in
dehydrated animals who did not drink. Average f during the 30 min after drinking
40.5

40.0
O
39.5

39.0

250
7
200
Et 150
-0
100
0

200
w _
- = 150
M I

u0m 100

50

0 15 30 45 60 0 30
Time (min)
Fig. 3. Effect of drinking water on rectal temperature (Tr), respiratory frequency (f) and
sweat rate of five dehydrated goats at Ta 40 'C. At 60 min, each animal was offered a
bucket of warm (36 °C) tap water and allowed to drink to satiation. The period of drinking
was 80+10 s and subsequent measurements began (time 0 after break on abscissa) at the
end of drinking for each animal.

NaCl was higher (P < 0 05) thanf during the last 30 min of dehydrated heat exposure
without drinking. There were no changes in Posm or Hct during the 30 min after
drinking saline, but PP measured at 30 min was lower than the pre-drinking value
(Table 1).
In two dehydrated animals which were allowed to look at a bucket of water for
5 min at the end of 90 min of heat exposure, the sight of water produced obvious
excitement but no sweating. At the end of the 5 min period, they were allowed to
drink and sweating occurred at 2-3 min after the start of drinking.
 RECOVERY OF SWEATING AFTER DRINKING IN GOATS 429
Effects of intraruminal water administration on thermoregulation
During the period of water deprivation preceding intraruminal administration of
water, the average loss in body weight was 4-0 + 0 4 kg and Posm, Hct and PP all
increased (Table 1). The time required to restrain the animal and to pass the tube

300 -

200

100
^

200
100

15 30 45 60 75 90
Time (min)
Fig. 4. Recovery of sweating after drinking in two dehydrated goats at Ta 40 °C. Top:
sweating after drinking water in a Nubian goat. Bottom: sweating after drinking saline
solution in an Alpine-Toggenberg goat. Each trace shows the output of a resistance
hygrometer in the effluent airstream from a ventilated skin capsule. The abscissa shows
the time after beginning measurements. There were no sweat peaks before 15 min. Arrows
mark the beginning and end of drinking.

through the mouth and into the rumen was approximately 30 s and the time required
to pump in the water was 87+8 s. In the four Alpine-Toggenberg goats, the
return of sweating was delayed compared to the return after drinking (Fig. 6) and the
latency to the first sweat peak was 13 min 42 s (range 9-22 min) after the beginning
of the pumping of water (Fig. 7, top). In these animals the change in sweat rate was
not statistically significant until the period 10-15 min after the water was pumped
in (P < 0 01) (Fig. 6). In the Nubian, however, sweating began even as the water was
being pumped in (Fig. 7, bottom). Because his response was different from the other
animals, and because the Nubian appeared to be more upset by the passing of the
stomach tube than the other animals, the experiment was repeated in this animal. In
the second experiment, he was restrained and the rumen tube was placed, but no
water was pumped in. The tube was left in for 3 min. He did not sweat. Then the tube
was removed and, after 10 min, placed in the rumen once more and water was
pumped in immediately. A sweat peak occurred 45 s after the end of pumping.
430 M. A. BAKER

DISCUSSION

In these experiments, dehydrated goats reduced cutaneous moisture loss to low

levels when they were exposed to heat. When they were allowed to drink, they began
to sweat within 2-3 min of the beginning of drinking. An osmotic signal acting on the

-
c

a)
-

CD
e

CDUE
C,,-.

0 15 30 45 60 0 30
Time (min)
Fig. 5. Effect of drinking saline solution on rectal temperature (Tr), breathing frequency
(f) and sweat rate of five dehydrated goats at Ta 40 'C. At 60 min, each animal was offered
a bucket of warm (36 °C) NaCl solution and allowed to drink to satiation. The period of
drinking was 147+6s and subsequent measurements began (time 0 after break on
abscissa) at the end of drinking for each animal.

brain via the blood or a signal initiated by vascular volume receptors cannot account
for the rapid onset of sweating after drinking water since sweating began before any
change in plasma osmolality or plasma volume was detectable. In dehydrated sheep
and goats, water may be retained in the rumen for a prolonged period after drinking
and plasma osmolality may change very slowly (Choshniak, Wittenberg, Rosenfeld
& Shkolnik, 1984; Blair-West, Gibson, Woods & Brook 1985). Osmoreceptors may
 RECOVERY OF SWEATING AFTER DRINKING IN GOATS 431
be present in the mouth, the stomach and the liver in some mammals. The evidence
for this has been reviewed recently by Bisset & Chowdry (1988). If osmoreceptors are
present in these regions in goats, stimulation of such receptors does not appear to be
necessary for the rapid onset of sweating after the beginning of drinking, since

40 5 -

4000 0000 00000

.00 .0.0

g.- 0.~
395 F
crV0-0
390 F
a I1t I -L

250
k& 4
200

-o4-ECu 150
a)
CD
100

50
I I"n a
-.I I I
oil -l -

150 -
CD-
a) _

-c 100 1
CuDM
(0 -
50

0 15 30 45 60 0 30
Time (min)
Fig. 6. Effect of intraruminal water administration on rectal temperature (Tr), breathing
frequency (f) and sweat rate of four dehydrated Alpine-Toggenberg goats at T. 40 'C. At
60 min, a tube was passed through the mouth into the rumen and water was pumped in.
The average time for passage of the tube and pumping was 117 s and subsequent
measurements began (time 0 after break on abscissa) when the tube was removed at the
end of the pumping period.

sweating occurs with the same latency whether the animals drink water or saline
adjusted to body fluid osmolality.
Dogs reduce the frequency of thermal panting when they are dehydrated and
exposed to heat (Baker, 1984). If they are allowed to drink, they pant at hydrated
levels within 1 min after the end of drinking (Baker & Turlejska, 1989). This rapid
recovery of panting is the same whether they drink water, when Posm falls
432 M. A. BAKER
significantly by 6 min, or saline solution, with no change in P.,,, and occurs before
there is a change in plasma volume. These experiments in dogs indicated that some
signals associated with the act of drinking, possibly from mechanoreceptors in the
mouth or pharynx, could be involved in the recovery of thermal panting to hydrated

200 -

150 F
100 F
50-

300

2000-
100

15 30 45 60 75 90
Time (min)
Fig. 7. Two different patterns of sweating after intraruminal water administration. Top:
late onset of sweating typical of four Alpine-Toggenberg goats. Bottom: early onset-of
sweating in Nubian goat. Each trace shows the output of resistance hygrometer in the
effluent airstream from a ventilated skin capsule. The abscissa shows the time after
beginning measurements. There were no sweat peaks before 15 min. Arrows mark the time
of insertion of the rumen tube and pumping of water.

levels, although the possibility that gastric distension might provide a stimulus for
the recovery of thermoregulatory evaporation was not tested.
In the present study, the experiments with a rumen tube were designed to find out
whether sweating might be initiated by the arrival of fluid in the rumen. The Nubian
sweated immediately when water was pumped into his rumen, indicating that the act
of drinking was not necessary for the rapid recovery of sweating. It would be
interesting to know whether either the act of drinking or the arrival of water in the
rumen would initiate sweating in this animal, but this would require further
experiments using a fistula to drain the fluid drunk before it reached the rumen.
In the four Alpine-Toggenberg goats, the act of drinking was apparently necessary
for the immediate recovery of sweating. Intraruminal water administration was
followed by sweating after a long delay, compared to the sweating which occurred
2-3 min after the start of drinking. It is difficult to imagine what receptors might be
involved in the late initiation of sweating after water was pumped into the rumen in
these animals. Ruminal stretch receptors or osmoreceptors would be expected to be
stimulated immediately upon the arrival of water. Hepatic osmoreceptors could be
stimulated with this time course even if water absorption was occurring slowly, but
 RECOVERY OF SWEATING AFTER DRINKING IN GOATS 433
the present experiments, do not shed any light on this possibility. Further studies
will be necessary to clarify both the individual differences in responses to
intraruminal water administration and the signals which might influence the
initiation of sweating after the manoeuvre.
The effect of dehydration on respiratory evaporation in these goats was not as
clear as its effect on sweating. Robertshaw & Dmi'el (1983) reported that dehydrated
Israeli black bedouin goats deprived of water for four days reduced the rate of
sweating but increased f and Eresp above hydrated levels when they were exposed to
high ambient temperatures. In the present study, f and Eresp were the same whether
the animals were hydrated or dehydrated and body temperature was higher in
dehydrated animals. Jessen & Feistkorn (1984) have shown that Eresp in goats is
driven by thermal signals from both the deep body and the brain. Brain temperature
was probably higher in the dehydrated animals than in the hydrated ones, since
brain temperature in animals with a carotid rete is controlled by the temperature of
arterial blood in the body core and by upper respiratory evaporation and blood flow
(Baker, 1982). Dehydration leads to a reduction in upper respiratory blood flow in
dogs (Baker, 1984) and may have the same effect in goats. Thus, the sensitivity of
the panting response to elevated internal body temperatures may have been reduced
in the dehydrated animals in this study. The increase in f above dehydrated levels
after drinking also supports this possibility; however, studies using controlled
stimulation of body core and brain thermoreceptors (Jessen & Feistkorn, 1984)
would be necessary to define conclusively the effects of dehydration on the thermal
responsiveness of panting in goats.
If the sweat rates of these goats are the same on the rest of the body as on the neck
where sweating was measured, then the rate of evaporative heat loss by sweating was
as high or higher than evaporation by panting in the hydrated animals. The peak
rates of Eresp measured in this study agree well with the rates reported by Jessen &
Feistkorn (1984) for resting Toggenberg goats with deep body temperatures around
40-41 TC, but sweat rates as high as those measured here have not been reported for
any domestic goat except for the black bedouin goat. Dmi'el et al. (1979) reported
that the mean sweat rate of 137 ml m-2 h-1 which they measured in black bedouin
goats exposed to solar radiation was about six times higher than that measured by
Jenkinson & Robertshaw (1971) in British Saanen goats exposed to an air
temperature of 40 'C. In another study, Dmi'el & Robertshaw (1983) measured sweat
rates of 144 g m-2 h-1 in bedouin goats in a climate climate chamber at a Ta of 46 'C
while animals exposed to solar radiation had higher sweat rates. In the hydrated
state, the Alpine-Toggenberg animals at 40 'C in the present study sweated at about
0 5 times the rate of bedouin goats at 46 'C and the Nubian at about 1-2 times this
rate, and sweat rates after drinking were usually higher. It seems likely that there is
a wide range of sweating capacities in different goat breeds and even among
individual animals which has not been appreciated because sweating has not been
measured in most studies of thermoregulation in goats in hot environments.
The act of drinking leads to a number of precipitous endocrine and cardiovascular
changes. Arterial blood pressure and central venous pressure rise acutely during
drinking (Hoffman, Phillips, Wilson & Schmid, 1977; Thrasher, Keil & Ramsay,
1987; Itoh, Oda, Asaeda, Sohma, Shigemi & Morimoto, 1988). In goats, the
434 M. A. BAKER
concentrations of adrenaline and noradrenaline in plasma rise during drinking
(K. Olsson & M. A. Baker, unpublished observations). Plasma levels of atrial
natriuretic factor rise (Januszewicz, Thibault, Gutkowska, Garcia, Mercure, Joli-
coeur, Genest & Cantin, 1986) and plasma arginine vasopressin (AVP) begins to
drop immediately after drinking (Thrasher et al. 1987; Thrasher, Nistal-Herrera,
Keil & Ramsay, 1981; Maltz, Olsson, Glick, Fyhrquist, Silanikove, Choshniak &
Shkolnik, 1984; Blair-West et al. 1985). The drop in plasma AVP occurs before
plasma osmolality changes and the mechanism is thought to be a neural one initiated
by receptors in the mouth stimulated by the drinking of fluid (Thrasher et al. 1987).
If the inhibition of sweating in dehydrated goats is effected by osmotic or other
stimuli acting on central neural regions controlling body temperature, then the rapid
onset of sweating after drinking may be initiated by neural signals from the mouth
or from the rumen acting centrally to overcome this inhibition. Changes in plasma
hormone levels or neural reflexes secondary to the cardiovascular changes during
drinking could also affect the secretion of sweat by acting centrally or peripherally
on sweat glands or cutaneous blood vessels. These possibilities would have to be
tested with further experiments.
This work was supported by National Science Foundation Grant DCB-8517709.

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