Comp. Biochem. Physiol., 1969, Vol. 31, pp. 347 to 363. Pergamon Press.

Printed in Great Britain

A COMPARATIVE STUDY OF THE TEMPORAL PATTERNS

OF CUTANEOUS WATER VAPOUR LOSS FROM SOME
DOMESTICATED MAMMALS WITH EPITRICHIAL
SWEAT GLANDS

T. E. A L L E N * and J. B L I G H

A.R.C. Institute of Animal Physiology, Babraham, Cambridge

(Received 17 February 1969)

Abstract--1. The temporal patterns of cutaneous water vapour loss from the
skin of the sheep, goat, ox, horse, donkey and llama during exposure to 40°C T a
and during i.v. injections and infusions of adrenaline are examined.
2. It is concluded that there are marked species variations in the extent to
which the discharge of sweat is affected by periodic contractions of the myo-
epithelium, or by simple displacement of fluid from the filled lumen as more fluid
is secreted into it.

INTRODUCTION
THE SWEAT glands of most mammalian species occur in association with the hair
follicles and have therefore been termed "epitrichial" (Bligh, 1967), but are
popularly classified as "apocrine". This latter term implies the breakdown of the
cell membranes and the extrusion of cellular contents in the formation of the
glandular secretion (Schiefferdecker, 1917), but it is doubtful whether the rate of
such a process could be varied to the extent necessary to produce the continuous
discharge of sweat which can occur in the ox during exposure to heat (McDowell
et al., 1954; McLean, 1963b) and in the horse during exercise (Evans,
1957). Sheep and goats, however, cannot sustain a continuous outflow of fluid
from their sweat glands, but discharge the glandular contents briefly and syn-
chronously from time to time (Bligh, 1961; Robertshaw, 1968). As with the
epitrichial glands of all species which have been examined histologically, those of
the sheep and goat are invested with myoepithelial structures (Jenkinson, personal
communication), and Bligh (1961) suggested that these discrete discharges are due
to the synchronous contractions of the myoepithelium, expelling the accumulated
products of a slow secretion which could well be apocrine.
It was further suggested by Bligh (1967), on the basis of available evidence of
sweat discharge patterns, that all epitrichial sweat glands may be derived from
glands with slow secretion and periodic emptying by myoepithelial contraction,
and that while in some species they may still function in this manner, other species
*Permanent address: C.S.I.R.O. McMaster Field Station, Mail Bag, Liverpool,
N.S.W., Australia.
i2 347
348 T . E . ALLEN AND J. BLIGH

may have evolved a capacity to increase greatly the rate of secretion in response to
an appropriate stimulus. In this circumstance a continuous discharge of fluid
could be by simple overflow of fluid from the filled lumen. The myoepithelium
may still contract from time to time causing brief variations in the rate of discharge,
or may have ceased to function.
Definite evidence of the involvement of two distinct processes in the discharge
of sweat in different species would require their experimental separation. Attempts
to inactivate myoepithelial functions while leaving secretion unimpaired have, as
yet, met with little success. Meanwhile, an examination has been made of the
patterns of aqueous discharge from the skin of the sheep, goat, ox, horse, donkey
and llama in circumstances which give some indication of the processes involved.
A raised ambient temperature was used as a natural stimulus. Single i.v. injections
of adrenaline, which simulate the brief discharges of sweat in sheep (Bligh, 1961),
were made to see if similar responses could be induced in the other species, while
continuous i.v. infusions of adrenaline were made to see if the adrenergic response
could be maintained and therefore attributable to an increase in the rate of sweat
formation.

MATERIALS AND METHODS

Animals
Details of the experimental animals are given in Table 1. The oxen, llama, horse and
donkeys were kept in loose boxes or small paddocks, while the sheep and goats were kept in
pens in a general-purpose animal house. There was no control over the environment to
which the animals were subjected before and between test periods in the climatic chamber
and no special instructions were given concerning feeding and management.

TABLE 1--DETAILS OF THE ANIMALSSTUDIED

Body wt.
Species Breed Identity Sex (kg)

Sheep (Ovis aries) Welsh Mountain N12 M (castrated) 45

Sheep (Ovis aries) Welsh Mountain x L3 M (castrated) 58
Clun Forest
Sheep (Ovis aries) Merino E9 M (castrated) 57
Sheep (Ovis aries) Soay* J 17 M (castrated) 26
Sheep (Ovis aries) Soay* K11 M (castrated) 25
Goat (Capra hircus) Sanaan 230 M (castrated) 91
Goat (Capra hircus) Sanaan 280 M (castrated) 68
Ox (Bos taurus) Jersey M 19 F 200-240
Ox (Bos taurus) Hereford x Shorthorn A40 M (castrated) 165
Horse (Equus caballus) Welsh Dainty F 240
Donkey (Equus asinus) Asiatic Gertrude F 130
Donkey (Equus asinus) Asiatic Dorothy F 137-169
Llama (Lama glama) -- Hyacinth F 80

* St. Kilda wild sheep (Jewell, 1961).

 PATTERNS OF SWEAT DISCHARGE 349

Measurement of loss of water vapour from the skin

T h e apparatus which was used to record changes in the humidity of the air as it passed
at a known rate over a known area of skin beneath a ventilated capsule was essentially the
same as that described by McLean (1963a). Air, drawn from a point within the climatic
chamber distant from the animal, passed through the ventilated capsule, which was sealed
onto the closely clipped skin with Durofix cellulose adhesive, and then through a spiral
copper tube which formed a double helix with that of the control stream of air. This served
to return the temperature of the air after it had passed over the skin to that of the control
stream. T h e air then flowed over fine (40 swg) copper-constantan dry-bulb and wet-bulb
thermocouples and through a flow-meter adjusted to maintain a flow rate of 1 1/min. A
control air stream, drawn from the same point in the climatic chamber, passed through a
system which was identical except for the ventilated capsule.
T h e wet-bulb and dry-bulb thermocouples of both air streams were connected to a
1 mV f.s.d. Cambridge multi-point recorder which completed each cycle of its eight input
channels every 48 sec. T o reduce the interval between the point recordings of the wet-bulb
temperature of the air stream which had passed over the skin, this thermocouple was con-
nected to each alternate channel of the recorder. This reduced the time interval between
items of information on this one parameter to 12 sec.
Ideally, when no water vapour was being added to the air as it passed through the
ventilated capsule, the two air streams had the same wet-bulb and dry-bulb temperatures.
When water vapour was being taken up from the skin the wet-bulb temperature of that
stream of air was raised proportionally, while its dry-bulb temperature, and both the wet-
bulb and dry-bulb temperatures of the control stream, remained unchanged. From this
information, together with the flow-rate and the area of the skin beneath the capsule, the
rate of water vapour loss could be calculated and conventionally expressed as g/m 2 per hr.
In practice it is difficult to standardize two wet-bulb thermocouples, and in order to
determine the extent to which deviations between them could be attributed to loss of water
vapour from the skin, the system was arranged so that during a check period the air stream
of the capsule line could be diverted around the capsule and through the remainder of the
system, while a third air stream passed through the capsule at the same rate and was then
discarded. This prevented the accumulation of moisture on the skin beneath the capsule
during the check on the wet-bulb temperatures. Any persistent difference between the
wet-bulb temperature at this time was taken into account in the calculation of the rates of
water vapour loss from the skin. T h e absence of leaks in the system was checked by momen-
tarily obstructing the air inlets and ensuring that the flow-meters then recorded zero flow.
T h e entire system was duplicated for the second ventilated capsule. Unless otherwise
indicated, the results given below refer to sweat capsules placed on the flanks just caudal to
the diaphragm on either side of the animals.
T h e system indicated the rate of uptake of water vapour by the air passing over an area
of skin. This was a measure of the rate of water vapour loss from the skin. T o avoid pre-
judgement of whether the water vapour was derived from passive physical diffusion through
the skin or discharge from the sweat glands, the non-committal term "water vapour loss" is
used in the presentation of the results.

Intravenous injections and infusions of adrenaline

A polyethylene cannula (0"1 m m i.d., 0.15 m m o.d.) was introduced into a jugular vein
of each animal a day or two previous to the sweat experiments. It was retained in position
by two skin sutures with 3-5 cm of the cannula projecting through the skin. T h e cannula
was kept filled with heparin-saline solution and plugged at its external end. T h e dead space
of the cannula was taken into account in all injections and infusions. Infusions were made
with a Palmer continuous-injection syringe pump. Adrenaline chloride solutions in isotonic
saline were used. All doses refer to the salt.
350 T. E. ALLEN AND J. BLIGH

Experimental procedure
During the first hour after the animal had been introduced into the climatic chamber the
air temperature remained steady or rose very slowly due to the presence of the animal. This
temperature varied from day to day between 20 and 25°C because it could not be made
completely independent of the building temperature when it fell within the same range. In
some experiments the ambient temperature (Ta) was kept between 20 and 25°C throughout
the experimental period. In others, at the end of the first hour the temperature of the
chamber was raised to 40°C during a period of approximately 45 rain and was then held
within + 1 °C for the remainder of the 7"5-hr experimental period.

RESULTS AND COMMENTS

Sheep and goats
Moderate (20-25°C) and high (40°C) ambient temperature ( Ta). The patterns of
cutaneous water vapour loss from all the sheep and the two goats tested were
essentially similar although there were differences between individuals, breeds and
species. No continuous loss of water vapour from the skin was detected in any
breed of sheep at either the moderate or the high Ta, but a sustained water vapour
loss of up to 50 g/m s per hr was usually recorded with the goats. This rate was
little affected by changes in ambient temperature. In both species brief discharges
of fluid occurred from time to time simultaneously beneath both capsules. The
highest rate of water vapour loss occurred within the first minute and then declined
during the following 3-8 min. These discharges were only occasional events in
sheep and very rare events in goats at moderate T a. They occurred much more
frequently in both species at the high T a when discharges might occur at intervals
of 5-10 min (Figs. 1, 2), but the frequency varied between individual animals, and

2.5oc

L_
1 I I I 1 I I I I . .I l ,,.I
Time, 5min intervals
FIG. 1. Spontaneous brief peaks in the rate of water loss from the skin of a Soay
sheep at 40°C T a. The peaks occurred synchronously beneath the ventilated
capsules on the right (R) and left (L) flanks of the animal. T h e discharges of fluid
from the skin are signalled by the abrupt changes in the wet-bulb temperature of
the air passing through the ventilated capsule, unaccompanied by any change in the
wet-bulb temperature of the air drawn directly from the room.
 PATTERNS OF SWEATDISCHARGE 351
even between periods of observation on the same animal. With one sheep, for
example, the average intervals between discharges at 40°C Ta were 11, 25 and 32 min
during different periods, although on another occasion only one discharge occurred
during 3.5 hr. Upon first exposure to 40°C Ta the discharges from the goat were

28 ¸

c
0
= 28
"o~ 26 2

~= Time, 5 rain intervals

FIG. 2. Spontaneous discharges of fluid beneath the ventilated capsule on either

side of the goat while T a was rising to 40°C. Successive discharges decreased in
magnitude and finally disappeared.

progressively smaller and were no longer detectable after the eighth or ninth
discharge (Fig. 2), but the response was better sustained during subsequent
exposure to heat, indicative of some acclimatization effect on the rate of sweat
secretion.
Single i.v. injections of adrenaline. The dose necessary to induce a discharge of
fluid varied between breeds in sheep. Whereas an i.v. injection of 0"4Fg/kg
elicited a discharge similar in size to the natural event in a Welsh Mountain sheep,
2-16 Fg/kg adrenaline was necessary to cause a discharge from a Soay sheep, and
this was considerably smaller than the natural event.
Although the first adrenaline-induced discharge of fluid from the skin of the
domesticated breeds of sheep at the moderate Ta was often equal in magnitude to
that of a single spontaneous discharge, repeated injections of adrenaline at intervals
of 15-30 min induced much smaller responses which could be only slightly en-
hanced by increasing the dose of injected adrenaline (Fig. 3). By contrast the
undomesticated Soay sheep did not show the same exhaustion phenomenon, but
even the largest doses of adrenaline produced an expulsion of only small quantities
of fluid which presumably did not empty the glands. The rise to the peak rate of
water vapour loss during the natural event was less steep than in the other breeds,
suggestive of a slower expulsion of fluid. That injected adrenaline does not
simulate the natural event in Soay sheep was emphasized by the occurrence of a
large spontaneous discharge of fluid shortly after an injection of adrenaline had
elicited only a feeble response (Fig. 4).
It was also difficult to induce discharges of fluid from the skin of goats with
i.v. adrenaline, 6/~g/kg being ineffective. However, by contrast with the Soay
sheep, the spontaneous discharges were also small.
352 T . E. ALLEN AND J. BLIGH

28
26
o
24 - D.B.
m"
22

~. 2o

i--- 18

20 20 20 40 I00
Fg A
I I I I I I I I I I I I I I I I
Time, 5 rain intervals

FIG. 3. T h e effect of i.v. injections of adrenaline upon the cutaneous water vapour
loss of a 45-kg Welsh Mountain sheep as indicated by changes in the wet-bulb
temperature of the air stream ventilating the capsule. After the initial discharge of
fluid in response to 20 Fg adrenaline subsequent injections of 20, 20 and 40/zg
adrenaline were much less effective. A slightly greater discharge occurred when
the 100 Fg of adrenaline was injected. T h e control wet-bulb temperature has been
omitted.

,-,~6
(:}.
QE 36

34 L b

~ 26

o ~

-~ E
~ 24

22

20
L
200 200 200 400 sp
I

$ /~g A
I I I I I I I I I
Time, 5 mln intervals

FIG. 4. T h e wet-bulb and dry-bulb temperatures of the air ventilating the

capsule (a) and the control stream (b) while single i.v. injections of adrenaline
were made into a 25-kg Soay sheep. Shortly after 400 Fg adrenaline had been no
more effective than three earlier injections of 200/zg adrenaline, a large spontaneous
(sp) discharge of fluid occurred.
 PATTERNS OF SWEAT DISCHARGE 353
Continuous i.v. infusion of adrenaline. Continuous i.v. infusions of adrenaline
into the sheep (up to 1"4/zg/kg per rain) and goats (up to 0.88/zg/kg per min) were
without effect.
Comment. Even in the absence of sweating some diffusion of water vapour
through the skin must occur. In sheep (Brook & Short, 1960) and humans (Kuno,
1934) with congenital absence of sweat glands this was found to be 12-31 g/m 2 per
hr and 11-33 g/m 2 per hr, respectively. A background water vapour loss was not
always detected, and it is possible that the instruments were insufficiently sensitive
for recording water vapour loss by simple diffusion, but this would not affect the
main conclusions which are based upon qualitative rather than quantitative
observations. Only rates of water vapour loss calculated to be greater than 50 g/m 2
per hr and influenced by at least one of the three stimuli are attributable to sweat
gland activity. When a continuous water vapour loss was recorded, it was little
affected by changes in ambient temperature or by intravenous injections or in-
fusions of adrenaline and is therefore attributed to transcutaneous diffusion.
The difference in the sweat discharge patterns between individual sheep at high
7", are not as great as those reported by Robertshaw (1968), but the number of
sheep was small and may be unrepresentative. The goat patterns are very similar
to those reported by Robertshaw (1968).
The decline in the size of the spontaneous discharges in the goat upon first
exposure to heat, and the decline in the size of the successive adrenaline-induced
discharges in the Welsh Mountain sheep are consistent with the proposition that
these discharges are by a myoepithelial contraction of pre-formed fluid. There is no
evidence that either the high T, or a continuous infusion of adrenaline can increase
the rate of secretion of fluid into the gland lumen sufficiently to cause a continuous
overflow of fluid, but some increase in sweat secretion must occur at high T, as
the size of the more frequent spontaneous discharge is maintained. This could be
the result of the local rise in skin temperature for when a shorn area of skin is
locally heated by i.r. lamps, the fluid discharged from this area in response to
repeated i.v. injections of adrenaline is greater than that from an unheated area of
the same animal (Allen, unpublished).
The spontaneous discharge is probably triggered by a neural rather than a
humoral stimulus as circulating adrenaline cannot simulate the natural event in the
Soay sheep. In the Welsh Mountain sheep, in which it can, there is no rise in the
systemic blood pressure at the time of the spontaneous discharge of sweat (Bligh,
1961). The breed difference in the effectiveness of i.v. injections of adrenaline
may depend upon the ease with which circulating adrenaline can penetrate to the
contractile elements.

OX

Moderate and high ambient temperatures. A low rate of water vapour loss from
the skin was recorded at 20-25°C T, immediately after the ventilated capsule had
been attached to the closely clipped skin. While T a remained steady there was no
change in this rate, but as soon as T a started to rise the synchronous stepwise
354 T.E. ALLEN AND J. BLIGH

increments in the rate of water vapour loss previously reported by M c L e a n (1963b)

were recorded (Fig. 5A). W h e n T~ was maintained at 40°C and the background
rate of continuous water vapour loss was constant at 100-200 g/m ~ per hr, brief
discharges similar to those recorded f r o m the sheep and goat were superimposed
upon it. T h e frequency of these peaks was approximately the same as that of the
stepwise increments at the onset of sweating.
183g/m z hr

20 5o ~oo 200 400

~g A

g 127 204 155

_~ C Baseline
F-t- q- - .
24 L 19 58 76 /.zg A/ram
I I I I I I I I I I I I I I I I ] I I I I I
Time, 5rain inlervols
FIG. 5. Effects u p o n c u t a n e o u s w a t e r v a p o u r loss f r o m t h e skin of a 200-kg Jersey
cow (A) when T a was raised to 40°C, (B) when adrenaline was injected i.v. and (C)
when adrenaline was infused i.v. In each section the upper trace is of the wet-
bulb temperature of the air that had passed through the capsule, and the lower one
is of the wet-bulb temperature of air drawn straight from the room. Differences
between these two wet-bulb temperatures are proportional to the rate of water
vapour loss from the skin as the dry-bulb temperatures of the two streams of air
remained identical. Figures above the traces indicate calculated rates of water
vapour loss (g/m s per hr). The vertical line in trace (A) indicates a 15-rain break in
the record. Figures below trace (B) indicate the size of doses of injected adrenaline
(Fg). Figures below trace (C) indicate rates of i.v. infusion of adrenaline (Fg/min).

Single i.v. injections of adrenaline. At the moderate T a i.v. injections of adrena-

line at dose levels of 0.13-0.3/~g/kg resulted in small brief discharges of fluid
which were similar to heat-induced discharges of the goat but not as large as those
of sheep. Larger doses of adrenaline (0.5-2.4/~g/kg) caused larger peaks which had
a more gradual rising phase t h a n were recorded from the goat and which also
declined m o r e slowly (Fig. 5B).
Continuous i.v. infusion of adrenaline. T h e continuous i.v. infusion of adrenaline
at the rate of 0.38 t~g/kg per min usually caused a rapid increment in the rate of
water v a p o u r loss, which was then sustained throughout the infusion period. At
other times this rate of infusion was ineffective. An increase in the infusion rate
f r o m 0.38 to 0.76/~g/kg per rain then caused the continuous discharge (Fig. 5C).
 VArTraNS or SWEATmscHAacr 355
Comments. McLean (1963b) confirmed several earlier reports that the con-
tinuous moisture loss from the skin of cattle exposed to a high Ta rises to 200 g/m ~
per hr or more, and showed that at the onset there is a series of stepwise increments
in the sweat rate which Bligh (1967) suggested may be due to periodic contractions
of the myoepithelium superimposed upon a rising rate of sweat secretion and
overflow. The new observation that these intermittent events continue as brief
peaks in the sweat discharge against an otherwise steady high rate of sweating is
consistent with this suggestion. However, these high rates of maintained water
vapour loss during exposure to heat, the increasing effectiveness of the progres-
sively increased doses of adrenaline and the continuous discharge of fluid elicited
glm 2 hr
o 30-- 43 95

~ 2 6 - --

~ 2o
~4 ~8
-- 16--1 I
Baseline
I ] I I l I I I I I I 1 I r [ I I 1 1 I ] I r 1
Time, 5 rain intervals
FIG. 6. Effect upon cutaneous water vapour loss from the skin of a 240-kg horse
of a rise in ambient temperature to 40°C. T h e u p p e r tracing is of the wet-bulb
temperature of air which had passed through the capsule. T h e lower tracing is
that of air drawn directly from the same point of origin within the chamber. T h e
second base-line check showed a slight error which was taken into account in
calculating the rates of water vapour loss indicated at two points.

by an i.v. infusion of adrenaline, all suggest that the principal effect of both heat
and adrenaline is to augment the rate of sweat secretion and overflow. Periodic
synchronous contractions of the myoepithelia still occur but exert only a transient
influence upon the rate of discharge. This interpretation is consistent with the
conclusions of Findlay & Jenkinson (1964), drawn from a histological study, that
the lumen of the bovine sweat gland does not empty when activated by heat, that
there is no apocrine process of secretion, and that the glands are functional
thermoregulatory organs.
Horse and donkey
Moderate and high ambient temperatures. Little or no water vapour loss was
recorded when the horse or donkey was at 20-25°C Ta, but when this was raised to
40°C there was a slow onset of a sustained water vapour loss from the skin (Fig. 6).
Rates of the order of 100 g/m z per hr were recorded from the horse, but the
simultaneous rates beneath the two capsules could be very different. On occasions
ofily very low rates of water vapour loss were recorded by either capsule system
356 T.E. A L L E N AND J. BLIGH

even when placed on areas which had previously yielded evidence of much higher
rates.
An unevenness in the rates of water vapour loss from the horse sometimes
occurred simultaneously beneath both capsules (Fig. 7), but these variations were
quite unlike the periodic discharges recorded from sheep, goat and ox. With the
C)

- 24-

~ 2 6 2 2

-- 2g

.Q
2 Is
i

d:l

1 I I I I t I 1 I I I I I I I I I I I
'rime, 5 rain intervals
Fxa. 7. Simultaneous fluctuations in the rate of water vapour loss f r o m the horse
as ambient temperature was being raised from 23 to 40°C. The ventilated capsules
were situated on the left (L) and right (R) flanks of the animal. The upper tracing
of each pair is the wet-bulb temperature of the air which had passed through the
capsule. The lower tracing is that of the air drawn directly from the room.

donkey the asymptotic rates of water vapour loss also varied with the position of
the sweat cups, the highest recorded rate being 166 g/m 2 per hr. The record was
smooth with no definite evidence of transient variations in the rate of water vapour
loss (Fig. 8A).
Single i.v. injections of adrenaline. Single i.v. injections of 1 ~g/kg adrenaline
caused an abrupt rise in the rate of water vapour loss from the skin beneath both
capsules in the horse and the donkey, and this was followed by a slow decrease to
the original rate. A smaller quantity of adrenaline (0.1-0.2 ~g/kg) caused a small
brief discharge pattern not unlike those elicited with much larger doses of adrenaline
in sheep. However, whereas in sheep progressively larger doses of adrenaline had
little or no effect upon the amount of fluid discharged, in the horse and donkey
increasing doses of adrenaline caused progressively greater responses, both in
magnitude and duration, with no evidence of exhaustion. Successions of injections
of the same amount of adrenaline were all equally effective (Figs. 8B, 9).
Continuous i.v. infusions of adrenaline. In both species the continuous infusion
of adrenaline at a rate of 0.1/xg/kg per rain in the horse and 0.25 ~g/kg per min in
the donkey caused the water vapour loss from the skin to rise rapidly to a rate of
about 200-300 g/m ~ per hr which was sustained during the period of infusion
(1-2 hr). Aqueous fluid was then dripping from the horse. This discharge of
fluid only slowly declined when the infusion was stopped, not ceasing completely
until 1-2 hr after the end of the infusion (Fig. 8C).
 PATTERNS OF SWEAT DISCHARGE 357

80 g/m z hr
30-
oo 28
26
2,4
~22
18

77 g/m a hr
~.- 28[-- B

I 2 5 IO 20 50 IOO /~g A
265 226 g/m z hr

20 ~ = 58p-g A/rain
] I I I I I I 1 1 I I I I I I I I I I I I I
Time, 5 rain intervals

FIG. 8. Effects upon cutaneous water vapour loss from the 130-kg donkey of (A)
a rise in ambient temperature to 40°C ; (B) i.v. injections of adrenaline and (C) an
i.v. infusion of adrenaline. Figures entered above the paired tracings of the wet-
bulb temperatures indicate the rate of water vapour loss (g/m 2 per hr). Figures
entered below the paired tracings indicate amounts of adrenaline injected i.v. in (B)
and rate of adrenaline infusion in (C).

10-22 IO-44 IO-57 I1-10 11-50 11-47 12-08

25 50 30 50 50 50 30
,Tr ~ ~
16

E
i ;: 12-27 '
35
12-41
40
'
12-53
50
'
15-28 hr
15-38
75
14-27
I00

2° I
E 14 I00 200 Fg A

(..) ....
16
14 I I 1 f I ] r I I
Time, 5 min intervals

FIG. 9. T h e effects of single i.v. injections of adrenaline into the 240-kg horse upon
the rate of water vapour loss from the skin. In each case the upper trace is that of
the wet-bulb temperature of air which has passed through the ventilated capsule,
while the lower trace is that of the wet-bulb temperature of air drawn directly from
the room. For each set of figures the u p p e r figure indicates the time of the i.v.
injection, while the lower figures indicate the a m o u n t of adrenaline injected (Fg).
358 T. E. ALLENAND J. BLIGH
Comment. The response to thermal stress and to injections and infusions of
adrenaline all indicate that the process of sweat formation can be greatly increased
and that expulsion need only involve the passive displacement of fluid from the
filled lumen as more is secreted into it. Even the rapid onset of an adrenaline-
induced sweat discharge cannot be attributed to myoepithelial contractions as it
can be greatly increased by increasing the quantity of injected adrenaline, and a
biphasic discharge pattern, which might be expected if the onset of discharge was
myoepithelial and the subsequent continuous outflow was simple overflow, was
never observed. This evidence supports the views of Takagi & Tagawa (1959)
that the principal process of secretion is not apocrine and that the myoepithelium
plays little or no part in the discharge of sweat in the horse and donkey.
No explanation can be offered for the fluctuations in the outflow of sweat
observed on some occasions only with the horse. They were quite unlike those
seen in the sheep, goat or cow, but were remarkably similar to those recorded when
the same apparatus was used to record cutaneous water vapour loss from a human
subject at high T a (Fig. 10). Such synchronous fluctuations in the output from

188 g/m 2 hr

~c ~.~ 24 26 ~
22
-~ ~ 24 o b
~ o-20
oE I I - 22 ~ S
-- 18 L o~

Time, 5 rain intervals

FIG. 10. S i m u l t a n e o u s fluctuations in the rate o f w a t e r v a p o u r loss f r o m beneath

v e n t i l a t e d capsules placed on the left ( L ) and r i g h t (R) sides o f the chest o f m a n
w h e n T a was raised to 40°C. T h e u p p e r tracing o f each pair is the w e t - b u l b
t e m p e r a t u r e o f the air w h i c h had passed t h r o u g h the capsule. T h e lower tracing
is that of the air d r a w n d i r e c t l y f r o m the r o o m .

the human atrichial sweat glands at different parts of the body have been reported
by Randall (1946), Albert & Palmes (1951), Nakayama & Takagi (1959) and
Custance (1962) although the frequency of the fluctuations varies considerably in
these different reports. Randall (1946) has attributed them to simultaneous
variations in the number of active sweat glands, while Nakayama & Takagi (1959)
have suggested that they could be due to periodic contractions of the myoepithelium.
The fluctuations in the sweat output of both the horse and man lack the features of
the brief natural discharges of sweat from the sheep and goat with the rapid rising
phase which can be readily, though not conclusively, attributed to myoepithelial
contraction, but this does not exclude the possibility of a slower contractile process
suggested by the peristaltic waves observed in the human atrichial glands by
 PATTERNS OF SWEAT DISCHARGE 359
Hurley & Shelley (1954). Whatever the correct explanation might be, there is now
the possibility that it is the same for the atrichial glands of man and the epitrichial
glands of both man and horse.

Llama
Moderate and high ambient temperatures. No water vapour loss from the skin,
either transitory or sustained, could be detected while the llama was at 20-25°C Ta.
When Ta was raised to 40°C a sustained water vapour loss became evident, and its
rate increased gradually and smoothly until it levelled out at between 100 and
250 g/m s per hr (Fig. 11A).

83 g/m 2 hr
52 - A
?
3O
28 _
_ Bose,,oe
26
E 24
oJ
_ g/m 2 hr
.d 22
6 22 44
.oi 20
eo -- B ,~-
II
-1 ii ~ l Baseline
I O0 Bose- 200 400
line
,u.gA
2°1-- C IlOg/m z hr
la! t
8
r 14
,
--I I I I t36 , /zg/min
, I , , I [ ~ I I I I I I
Time, 5min inlervals

FIG. 11. Effects upon cutaneous water vapour loss from the 80-kg llama of (A) a rise
in ambient temperature to 40°C; (B) i.v. injections of adrenaline and (C) an i.v.
infusion of adrenaline. Figures entered above the paired tracings of w e t - b u l b
temperatures indicate the rate of water vapour loss (g/m z per hr). Figures entered
below the paired tracings indicate the amounts of adrenaline injected i.v. in (B) and
rate of adrenaline infusion in (C).

Single i.v. injections of adrenaline. Intravenous injections of adrenaline at dose

levels of 1.25-2-5 Fg/kg induced patterns of water vapour loss very similar to those
observed in response to the high ambient temperature. There was increase to a
steady value followed by a slow decline. Increasing the amount of injected
adrenaline caused some increase in the asymptotic rate of water vapour loss and
markedly extended the duration of the discharge (Fig. l lB).
Continuous i.v. infusions of adrenaline. Infusions of adrenaline at a rate of
0.25/zg/kg per min resulted in a gradual rise in the rate of water vapour loss to
110 g/m s per hr, but the rate of discharge was not maintained, and declined
before the infusion was stopped (Fig. 11C). An increased rate of adrenaline infusion
360 T. E. ALLENANDJ. BLIGH
(0-5/xg/kg per min) caused the rate of loss to rise to 140 g/m 2 per hr; this was
maintained until the infusion was stopped and then declined only slowly.
Comment. There are no features of the sweat gland discharge patterns, in
response to heat or circulating adrenaline, which could be attributed to myoepi-
thelial activity. The response to all three stimuli suggests that the rate of secretion
is increased and that discharge of fluid onto the skin is the result of simple overflow.
However, unlike the horse and donkey in which the secretory processes are quickly
brought to a high level of activity by the adrenergic stimuli, those of the llama are
recruited more gradually. As the onset of sweating in response to heat is also
gradual, it is probable that this pattern relates to the properties of the secretory
cells rather than to a slow diffusion of adrenaline to the secretory cells.

DISCUSSION
Evans (1957) commented that not only are there considerable differences
in physiological properties and functions between the epitrichial and atrichial
glands which make generalization misleading, but also that even within each type
there is not complete uniformity, while Bligh (1967) pointed out that there is no
clear indication of the extent of the functional differences that might exist between
histologically similar epitrichial glands of different species.
The purpose of the experiments reported here has been to remedy this deficiency
by comparing the sweat discharge patterns of several mammalian species with
epitrichial sweat gland in identical experimental circumstances, and considering
whether the results are explainable in terms of secretory and myoepithelial
activities.
Differences in the temporal patterns of sweat discharge in response to thermal
stress may vary considerably between individuals as well as between species
(Robertshaw, 1968) and the sweat patterns reported here, which have been obtained
from only one or two animals from each species or breed, may not be representative,
but the between-species differences are great compared with the within-species
differences and the patterns reported here are to some extent confirmatory of
earlier reports (see Bligh, 1967). There are some new observations: the peaks
superimposed upon a steady rate of sweating in the ox, the irregular but synchron-
ous variations in the outflow of sweat from the horse, and the slow onset and
completely smooth sustained discharge of fluid in the llama.
The evidence reported here confirms that the epitrichial glands of different
species vary greatly in the rate at which sweat is formed in different circumstances,
and in the pattern of its discharge onto the skin. The sheep seems to rely entirely
upon the myoepithelium to expel the glandular contents which are not secreted
fast enough to cause a continuous passive overflow onto the skin in response either
to heat or to circulating adrenaline. In complete contrast, no evidence was obtained
of myoepithelial activity in the sweat glands of the llama. Both a rise in ambient
temperature and an increase in the level of circulating adrenaline apparently
caused a slow increase of the rate of secretion to a new steady rate at which fluid
w a s continually being displaced from the lumen onto the skin surface. The ox is
 PATTERNS OF SWEAT D I S C H A R G E 361
intermediate between the sheep and llama with a continuous overflow of sweat
onto the skin when the rate of secretion increases in response to either heat or
intravenous adrenaline, but with periodic peaks in the sweat outflow during
exposure to heat attributable to myoepithelial activity. This might be cited as an
example of a species in which myoepithelial activity is no longer of any functional
significance, but still occurs. The horse and the donkey largely parallel the llama
as species with a capacity to secrete at a high rate, so that sweat discharge
is by simple overflow. Indeed, the rate at which sweat secretion is recruited
in response to an intravenous infusion of adrenaline in these species is much
greater than in the llama. The simultaneous fluctuations in the outflow of sweat
which were sometimes recorded from the horse were very similar to the variations
in the outflow of sweat from the atrichial glands of man and which could be due to
myoepithelial activity.
Although these patterns of sweat discharge could be placed in an order which
complies with the suggestion by Bligh (1967) that the glands of modern mam-
malian species may all have derived from glands with slow rates of secretion and
intermittent expulsion, this is not evidence for the hypothesis. However, even if
the myoepithelial structures which surround the atrichial glands are quite inactive
as Sperling & Koppanyi (1949) have surmised those of the cat's pad to be from
their histological appearance, their very presence might suggest a common origin
with those of the epitrichial sweat glands and the mammary glands (Hyman, 1942).
Within each anatomical classification there may be glands which vary in functional
details, with some overlap between the two groups. The atrichial glands of the
cat's pad have been used to further the understanding of the human atrichial
glands of man, and marked similarities in both innervation and secretory processes
have emerged (Lloyd, 1959), but there may be no more validity for the assumption
that those of the one species can be regarded as the experimental model of those of
the other than there would now be for regarding the epitrichial glands of the sheep
as being essentially the same as those of the ox, horse or llama.

SUMMARY
1. A ventilated capsule technique has been used to record the temporal patterns
of cutaneous water vapour loss (sweat discharge) from the sheep, goat, ox, horse,
donkey and llama in response to (a) a rise in ambient temperature (Ta) from
20-25 to 40°C; (b) i.v. injections of adrenaline and (c) i.v. infusions of adrenaline.
2. The sweat glands of the sheep and goat did not discharge continuously in
response to either heat or injected adrenaline, but discharged spontaneously
briefly and synchronously over most, if not all, of the skin surface from time to
time. This event occurred more frequently when Ta was raised. Similar dis-
charges could be induced by i.v. injections of adrenaline, but large doses were
sometimes necessary with both species. The discharge is attributed to myoepi-
thelial contraction. The spontaneous event is probably under the control of the
sympathetic nervous system.
362 T . E . ALLEN AND J. BLIGH

3. T h e ox responded to heat and to infused adrenaline with a continuous

output of sweat. T h e stepwise onset of sweating in response to heat and the
regularly occurring brief increments on the steady flow of secretion were indicative
of periodic contractions of the myoepithelium in addition to the secretory response.
Adrenaline injected or infused i.v. had a prolonged effect suggesting a stimulatory
action u p o n sweat formation.
4. T h e horse and donkey responded to b o t h heat and infused adrenaline with
a continuous discharge of sweat, higher rates resulting f r o m i.v. adrenaline than
f r o m a raised T a. Sometimes the discharge of sweat was smooth, but on other
occasions synchronous irregular fluctuations in the discharge rate were observed.
T h e s e lacked the characteristics of myoepithelial involvement and are unexplained.
5. T h e llama responded to both heat and to adrenaline with a slow smooth
onset of sweating. N o fluctuations in outflow suggestive of myoepithelial involve-
m e n t were seen.
6. T h e s e differences in the patterns of sweat discharge f r o m the epitrichial
glands of m o d e r n species give some support to the proposition that those glands
which can maintain a continuous output of sweat in response to heat may have
evolved f r o m glands which could secrete sweat only slowly and in which discharge
was affected by periodical contraction of the myoepithelium.
Acknowledgements--We are most grateful to Mr. A. J. Barton for his technical skill in
assembling and maintaining the equipment, and to Messrs M. Beadle and S. Mills for
their enthusiastic help in the conduct of the experiments.
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Key Word Index--Sweat formation; sweat discharge patterns; atrichial sweat glands;
epitrichial sweat glands; apocrine sweat secretion; eccrine sweat secretion; myoepithelium;
adrenaline, effects on sweating; sheep sweat pattern; goat sweat pattern; ox sweat pattern;
horse sweat pattern ; donkey sweat pattern; llama sweat pattern.