Tica JOURNAL

Copyright

PHARMACOLOGY

1967

by

THE

The

AND

SYMPATHETIC

NERVOUS

MECHANISM.

II.

ON

THERAPEUTICS

EXPZRIMENTAL

EFFECT

BODY

R. P.
and

N.

HORMONES

MAINTENANCE

STERN,

157, No.
in U.S.A.

HOMEOSTATIC
OF

ADRENALECTOMIZED
D.

of Chemical
Departments

ADRENOCORTICAL

TEMPERATURE

MAICKEL,

AS

SYSTEM

OF

COLD-EXPOSED

Laboratory

Vol.
Printed

& WilkinsCo.

Williams

RATS

E. TAKABATAKE4

B. B. BRODIE

AND

Pharmacology,
National
Heart
Institute,
Bethesda,
Maryland
of Pharmacology
and Psychology,
Indiana
University,
Bloomington,
Indiana
Accepted

for

publication

February

16, 1967

ABSTRACT

MAICKEL,
R. P., D. N. STERN,
E. TAKABATAKE
AND B. B. BRODIE : The
sympathetic
nervous
system
as a homeostatic
mechanism.
II. Effect of adrenocortical
hormones
on body temperature maintenance
of cold-exposed
adrenalectomized
rats. J. Pharmacol.
Exp. Therap.
157:
111-116,
1967. Intact
rats exposed
to low environmental
temperature
(4#{176}C)maintain
body
temperature
by piloerection,
vasoconstriction,
shivering
and mobilization
of glucose
and
free fatty
acids. In contrast,
totally
adrenalectomized
rats are unable
to activate
any of
these
systems
; they
lose body
heat rapidly
and die in a few hours
when
exposed
to low
temperature.
The sugarand lipid-mobilizing
actions
of exogenous
catecholamines
are also
markedly
reduced
by adrenalectomy.
The inability
of adrenalectomized
rats to survive
in
the cold can be reversed
by acute
treatment
with glucocorticoids
such as cortisone,
dexamethasone
or prednisolone
or by chronic
treatment
with aldosterone.
This treatment
also
restores
responsiveness
to catecholamines,
suggesting
that the adrenalectomized
rat may be
considered

as

of effector

systems

The
has

preceding

a functionally

paper

described

the

sympathectomized

which

normally

(Maickel

similarity

et
of

that

mals

to

the

failure

maintain

of

adrenalectomy

ministration

of

istration

of

sonable
due
A
cate

were

cortisone,

the

defect

to the lack of adrenal

number
of reports
adrenalectomized

ani-

reversed
not
it

in

in

by
by

seemed

adrenalectomy

cortical
in the

1960
the

sympathectoevidence
sugtemperature

but

hormones.
literature

animals

are

sympathetic

to

1939 ; Sawyer

for

publication
in part

October
by U.S.

cold-exposure

In both
of
adrenalectomized

ad-

administration

rea-

in

was

be due

these
of

cold-exposed,
to the

lack

poorly

21, 1966.
Public
Health

to

to

test

reverse

also

adrenalectomized
responses
to

Grant
MH-06997.
Present
address
: Departments
of Pharmacology
Psychology,
Indiana
University,
Bloomington,
47401.
5 Present
address:
Department
of Psychiatry,
New York State
Psychiatric
Institute,
New York,
N.Y.
4 Present
address
: Pharmaceutical
Faculty,
Nagasaki University,
Nagasaki,
Japan.
Service
2

and
Ind.

effects
used
sistent
cortical

of
as

or

responsiveness
111

regulation
function.

of

ability

of

of

would

the
impaired
mechanisms
in
rats
was
de-

various

steroids

adrenalectomy.

animals
sugar-

also
and

peripheral

Since

show
decreased
lipid-mobilizing

these
The

of
by

hydrocortisone.

of adrenocortical

hypothesis
maintain
of

1933).

rats

comparison.

with
the
hormones

known

adrenalectomized

catecholamines,
a

at.,

the
responses
can
be restored

of temperature

effects
the

well

Schlossberg,

cortisone

the

the

et

1963) . Similarly,
adrenalectomized

The
present
investigation
response
of energy-producing
cold-exposed,
adrenalectomized

mdi-

Shafrir

is
and

situations,
animals

breakdown

catecholamines

1960;

and
Brodie,
sensitivity
of

(Secker,

the

of

Steinberg,

animals

Thus,

responsiveness

administration

; Maickel
increased

signed
Received
1 Supported

impaired

function.

and

of
the
admin-

of

to

(Shafrir

in

due to a breakdown
functions.
Since

catecholamines,

that

that

body

because

by

responsive

1967)

of adrenalectomized

normal

a cold enviroment
was
peripheral
sympathetic
effects

at.,

animal

activated

responses

adrenalectomized
and
chemically
mized
rats
exposed
to cold.
The
gested

are

responses
results
that

the
the

were
are

con-

adrenofunctional

sympathetic

ef-

112

ET

MAICK.EL

fectors.

Thus,

may

also

the

be

adrenalectomized

considered

as

animal

Adult,
male,
Sprague-Dawley
g) were used in all experiments.
rats
were
obtained
from
the

(170-200
ectomized

Assay

Laboratories,

Chicago,

Ill., and

were

of

estimated

mean

survival

main-

time

(M.S.T.)
and
rate
of change
of heat
content
(-dQ/dt)
were
made as described
in the preceding
paper
(Maickel
et al., 1967).
Materials.
Cortisone
acetate,
aldosterone,
dexamethasone,
prednisolone,
d,t-norepinephrine
hydrochioride
and
t-epinephrine
bitartrate
were
commercially
available
compounds.
RESULTS.

Effect

of NaC1

deprivation

an

impaired

was
take

made
on

mals

to

of partial

salt

deprivation
Rectal

Days

to

maintain
each
the

body

of

tap
of

reduced

was

the
their

given

rats

drinking
the greater

ability

table

of

2,

glucose
cold.
It

not prevent
while
NaCL

rats

higher

supplied

by

used

up-by

perature

level

Reversal
of
acute
treatment

glucose.

was

36.7
36.6
35.0
35.6

0.8
0.5
0.5
0.6

34.4
33.9
31.1
30.0

31.7
32.2
27.8
25.6

of adrenalectomized

Cold-Exposure

in
did

to the
anithe animals
heat
in
owing

The

energy

was
the

falling

rapidly

body

tem-

rapidly.
by
dep-

rats

exposed

.-dQ/dt

4hr
hr

28.3
28.9
21.1
20.6

to cold

for
3hr

2.9
0.6
2.5
2.5

or

2hr

2.2
0.5
1.6
1.3

NaCl

of
adrenalectomy
steroids.
Since
Na

oc

0
1
2
3

ability

temperature,
as the NaC1-

M.S.T.
lhr

is

the

lose much
presumably

exposure

on H,O
Obr

mainexposed

1%

however,

animals
effects
with

to

conclusion

provided
note
that

of blood
of

of

NaC1-5%
imbalance

of

loss in body
as effective

source,

4 hr
of these

rate

rate

when

did not
exposure,

this

rate.

the

body
temperature
that
glucose
alone

glucose
mixture
usually
mals.
It is interesting
to

to the

rapid
the

drinking

to maintain
is evident

With
intake,

animals

comparison

the rapid
alone
was

cold.

1%
Na

this

the

3 days
rats
to

Na

twice

the

for
a

of adrenalectomized
5%
the

1, 2 or

temperature

evidence
in

when

water,

than

inani-

1 show

a more

tap

more

internal

to cold.
Further

at

drinking

study
Na

table

decreased

fell

of

has

in the

of

temperature
loss

in

water
for
adrenalectomized

day

adrenalectomized
glucose.
Apparently,
tam

Nat,

temperature

data

temperature

3 days
heat

animal

retain

body

The

body

After

to

their

cold.

succeeding

temperature
after

Temperature

maintain

given
only
glucose
the first
2 hr of

on body

on body

ability

157

adrenalectomized

adrenalectomized

effects
of drinking
on
the
ability

TABLE

Eff!ts

cold-exposed

the

of the effects
of a decreased
the
ability
of adrenalectomized

exposed

tamed
on normal
laboratory
chow
(containing
0.5% NaCl)
and
drinking
water
containing
1%
NaCl-5%
glucose
until
used
4 to 10 days
after
surgery.
Measurement
of plasma
levels
of corticosterone
by the
method
of Guillemin
et at.
(1959)
indicated
no measurable
steroid
content.
Animals
were
exposed
to cold
(4#{176}C)in individual
screen
wire cages. The steroids
used were
commercially
obtainable
materials,
administered
as described
in the tables.
Rectal
temperatures
were
measured
with
a
small
animal
probe
and a Yellow
Springs
TeleThermometer.
Animals
were stunned,
decapitated
and
blood
was
collected
into
heparin-treated
beakers.
After
transfer
to tubes,
the blood
was
centrifuged
immediately
and aliquots
of plasma
were
used
for the
determination
of free
fatty
acids
(FFA),
by the
method
of Dole
(1956).
Blood
glucose
levels
were
determined
with
the
Glucostat
(Worthington
Biochemical
Corporation)
reagent,
and liver glycogen
was determined
by the method
of Carroll
et at. (1956).
Calculations

of

Since

rats.

rats
AdrenalHormone

METHODS.

Vol.

temperature

functionally

animal.

sympathectomized

AL.

3.4
1.2
3.2
2.3

24.5
23.9
18.9
12.8

2.3
2.0
3.4
1.4

7.9
7.2
5.2
3.8

cal/kr

546
603
765
1083

Each
value
is the mean
S.E.
of 14 animals.
Animals
were maintained
on 1% NaCl-5%
glucose
in
drinking
water
for 4 days after surgery,
then tap water
was substituted
for 1, 2 or 3 days prior
to coldexposure.
The fall in body temperature
of animals
maintained
on NaC1-glucose
did not vary significantly
when exposed
to cold on days 4 through
10 after surgery.

1967

HOMEOSTATIC

MECHANISMS.

TABLE

Differential

effect

of drinking

water
maintain

containing
NaCI
body temperature
Rectal

113

II

or glucose
on ability
of adrenalectomized
when exposed
to cold

Temperature

after

rats

for

Cold-Exposure

Treatment

M.S.T.
Ohr

3hz

2hz

lhr

Each value
is the
mals were maintained
described
below.

36.7

0.8

34.4

36.7
36.9
36.1
36.6

0.4
1.2
0.7
0.6

32.8
33.4
32.2
31.7

hr

cal/hr

2.1

31.7

2.9

28.3

3.4

24.5

2.3

7.9

546

0.6
0.4
0.5
1.1

30.6
32.2
31.1
28.4

1.4
0.6
1.0
1.9

28.9
30.0
25.6
27.2

1.4
0.3
1.8
1.8

27.2
27.7
18.3
18.9

1.5
1.1
2.0
2.6

8.4
8.6
6.0
6.1

461
437
846
841

mean S.D. of six animals

on 1% NaCl-5%
glucose

except
that
for 4 days

effect

of acute

administration

of various
exposed

Rectal

NaCl-glucose
after
surgery,

steroids
to cold

controls
are
then
groups

on adrenalectomized,

after Cold-Exposure

Temperature

14 animals.
Aniwere treated
as

TABLE

Protective

salt-deprived

M.S.T.
Ohr

lhr

2hr

3hr

35.0
36.1
35.6
36.0
35.6

Control
Aidosterone
Cortisone
Dexamethasone
Prednisolone

0.5
0.4
0.6
0.6
0.8

31.1
31.7
84.5
82.8
32.2

1.6
1.3
0.4
0.4
0.5

27.8
28.9
83.4
8L2
81.7

2.5
2.4

0.5

0.8
1.0

21.1
23.9
32.8
SI.
30.0

seemed

to

be

to

the

effect
of various
steroids
administered
cold-exposure
was examined.
The
test

studies
mized

glucose

adrenalectomized
tap
water
for

of steroid

was

administered

exposure.

The

rectal

subsequent
that

the

prior

to

(table
except

reversing

the

3)

heat
would
verse

of the
adrenalectomized
rats.
Thus,
it
seem
that
aldosterone
was unable
to rethe
effects
of adrenalectomy
and
saline
period

of treatment.

1.8

H20 controls
are 13 animals.
Animals
acetate
(26 mg/kg
i.p., as an aqueous
(0.126 mg/kg
i.m., as an aqueous
soluto cold. Values
in italics
are signifi.05).
of

effects

of

administration
were

animals

carried
were

adrenalectomy

of

by

steroids.

out in which
given
daily

Further

adrenalectodoses
of the

of adrenalectomy
with
table
3, these

effective

in a 4-hr

30.8

765
684
186
209
238

steroids

demonof

1.1
-

cal/kr

5.2
6.0
21.7
18.2
18.3

dur-

aldosterone
loss

81.7

3.4
4.9

dose

changes

steroids

cold-

1%

single

4 hr

of

were

deprivation

in

days.

1.8

18.9
21.7

hr

steroids
during
the
2-day
period
while
they
were
drinking
tap water
instead
of 1% NaC15% glucose.
Under
these
conditions,
all of the

that

rats
instead

temperature

cold-exposure
all

the

Reversal

tem-

chronic

NaC1-5%

ing

to

pro-

system
chosen
was
had
been
drinking

strate

damaging
mechanisms,

3.2
3.4
0.8
0.5

Each
value
is the mean
S.D.
of nine animals,
except
that
were maintained
on tap water
for 2 days,
then given
cortisone
suspension),
prednisolone
(5 mg/kg
s.c., in oil), dexamethasone
tion)
or aldosterone
(1.25 mg/kg
s.c., in oil), 4 hr before
exposure
cantly
different
from untreated
controls
at each time point
(P <

perature-maintaining

-.dQ/d:

4hr

oc

tective
prior

rats

for

Steroid Treatment

rivation

-dQ/dt

4hz

.c
1% NaCI
5% glucose
1% NaCl
(1 clay)
1% NaCl (3 days)
5% Glucose
(1 day)
5% Glucose
(3 days)

to

body

tion

tested

of

aldosterone

adrenalectomy
cortisone,
gesting
tion.
Effect

were

able

to

(table
results
in

reversing

is a slower
dexamethasone
different
locus

a
of

steroid

reverse

the

effects

4).
When
compared
suggest
that
the acthe

process

effects

than

or prednisolone,
or mechanism

hormones

on

lipolytic

that

of
of

sugof ac-

and

ET

MAICKEL

TABLE
Protective

effect

of chronic

administration

Temperature

after

on adrenalectomized,

Cold-Exposure

salt-deprived

M.S.T.
lhr

2hr

3hr

Control
Aldosterone
Cortisone
Dexamethasone
Prednisolone

35.0
36.1
36.2
36.7
36.1

0.5
0.6
0.4
0.4
0.6

31.1
35.0
86.1
37.2
35.0

1.6 27.8
0.8 33.9
0.9 35.0
0.4 35.6
0.9 53.4

2.5
1.4
1.5
0.4
1.1

21.1
31.7
33.9
53.9
32.5

3.2
18.9
5.4 82.2
1.4 31.1
0.8
1.3 30.6

Each
value
is the mean
S.D.
of seven
animals,
except
that
H,O controls
were maintained
on tap water
for 2 days,
and steroids
were given once daily
table
4. The exposure
to cold was made
approximately
16 hr after
the last
italics
are significantly
different
from controls
at each time point
(P < .05).
TABLE
of steroid

pretreatment

on glycolytic

and

3.4
1.1
1.4
2.1

hr

cal/hr

5.2

765

23.2

185

19.3
20.5
19.5

242
177
262

are 13 animals.
Animals
at the doses described
in
dose of steroid.
Values
in

lipolytic

response

to catecholamines

FFA

Plasma

in adrenalectomized
Plasma

Glucose

Control

Norepinepbrine

Control

pEq/ml

Intact control
Adrenalectomy
+
1%
Adrenalectomy
+
tap water
(2 days)
Acute
Aldosterone
Cortisone
Dexamethasone
Prednisolone
Chronic
Aldosterone
Cortisone
Dexamethasone
Prednisolone

0.35

0.05

0.37

mg/I

0.03

0.07
0. 10

112
87

3
5

0.45

0.08

0.41

0.04

59

0.27
0.33
0.37

0.01
0.06
0.06

0.31
0.71
0.57

0.04
0.11
0.15

81
86

0.34
0.40
0.36

0.03
0.03
0.06

0.64 d
0.80
0.87

00 ml

mg/g

52
ii

14
16

31

io

6
2

78

15

89

10
11

12
23

4
6

117

138

16
5

136
17

95

0.08

77
113

0.06

0.16
107

(Control)

Epinephrine

0.84
0.49

rats
Uver
Glycogen

Treatment

Animals

-dQ/di

4hr

oc

Effect

rats

for

Treatment

Ohr

157

of various
steroids
exposed
to cold

Rectal
Steroid

Vol.

AL.

ii
12
15

17

29

11

195

10

Each
value
is the mean
S.D. of six animals.
Steroid
pretreatments
were as described
in table
3
(acute)
and table
4 (chronic).
Plasma
FFA elevations
were induced
by norepinephrine
(0.1 mg/kg,
aqueous,
i.m.) and blood glucose
by epinephrine
(0.2 mg/kg,
in oil, i.m.),
30 mm before
measurement
of
blood levels.
Values
in italics
indicate
significant
response
to catecholamines
(P < .05).
glycolytic

actions

ectomized
plasma

animals.
FFA
and

ogenous

of

catecholamines

of

in

adrenal-

The
increased
levels
blood
glucose
evoked
by

in adrenalectomized
1967) . This
similarity
responses

catecholamines

were

markedly

animals
(Maickel
between
the

adrenalectomized

animals

exposure
of
ex-

and

cholamine
results
of

reduced

treatment

et at.,
reduced

tions
rats

to

plasma

cold-

of
are

to

seemed
studies
on

the

the

administration

lipolytic

catecholamines
shown
in table
FFA

levels

of

more
than
coincidental.
of the effects
of steroid

to

and
in
5.

cateThe
pre-

glycolytic

adrenalectomized
The
response

norepinephrine

mirrored

acof

1967

HOMEOSTATIC
results

obtained

with

administration

the

of

cortisone

restored
nephrine,
In

or

chronic

steroids

the

results

somewhat
general

blood

was

dose

animal

aldosterone

had

no

of

glycogen.
the results

This
possibility
obtained
after

oids

for

days.

sumably

Again,
had
a

reflecting

activity

of this

the

lack

of

adrenal

opinion
corticoids

causes

stress,

but

tomized

animal

can

survive

trolled
Ramey

environment.
and
Goldstein

the
data
tent

presented

steroids
tional
enabling
tation
The
tical
rats

steroids

adequate

paper

that

the

consis-

to

maintain
the
funcof the
effector
systems,
to respond
to changes
in
which

require

adap-

of

In

kept

to
fact,

alive

intake

of
to

the

for

saline
situations

adrenal

some

cor-

extent

by

adrenalectomized
some

time

provided
requiring

on
they

These

animals

extra

energy

substrates

heat.

The

1000

rate

of

exacerbates
ciency

the
and

lvtes

be
of

of

that

in

situations

was

rate

insuffi-

of

electro-

way

for

to

increased

energy

particularly

pertinent

animals,
exposed
saline,
display
no

to cold
signs
of

vasoconstriction

or

Furthermore,

even

catecholamines

do

not

increase

responsive

to

changes

are
of

that
glucose

exogenous

levels

that

of fatty

effector

responsiveness
by

adrenalectomized
are
unable

temperature

the
to

no

longer

That

involved

suggested

or glucose,

are

catecholamines.

the

shivering.

acids

systems

probably

the

respond

piloerection,

suggesting

to

loss

sympatheclack of saline

some

an

seems

adrenalectomized
2 days without

heat

rats

requiring
It

un-

and

corticoid

adrenalectomized

sur-

were

imbalance

responsible

of

cold

(cal/br)

effects

suggest

may

that
after

calories/hr

tempera-

the

4 hr.

to that
seen after
chemical
The results
show
that
the

in

ionic

this

break-

to

catecholamines

studies

which

show

rats
maintained
maintain
their

on
body

in the cold.

Evidence

that

the

effects

of

adrenalectomy

are due to
replacement

the lack of steroids

comes
from
the
experiments
described
in tables
3

and

example,

4. For

pretreatment

of adrenalec-

tomized
rats
with
or prednisolone
4
enables
the
animals

cortisone,
dexamethasone
hr prior
to cold-exposure
to maintain
their
body

temperature.

steroid-pretreated

The

had

-dQ/dt

than
rats.

%
In

that
of
contrast,
aldosterone

dose

of

an

tive

action

are

mals

lost

extra

in

than

of

adrenocortical

placed

in

deprived

more

is further

The

are

than
body

down

action

obscure.

present

is replaceable

subjected

increased
adrenalec-

of

be

action

saline.
be

the

systems.

supportive

can

that

1957,
the

conditions

ordinary

well

As
recently
as
(1957)
considered
to

are
necessary
responsiveness
the animal
of such

is

con-

hypothesis

environmental

not

in

site

in the

the

the

of
ab-

animals

similar
tomy.

to

careful

hormones

with

an

that

physicochemical
adrenal

ability
withstand

the
and

to
succes-

cold-exposure

finally

less

the

exposure
each

deprivation,
decline

to mobilize

longer

until

rapid

3 days

expenditure.

treated
pre-

held

able

saline

more

for

inability

glucocorticoid

has

to

primary

of

returned

conditions

general

sensitivity

by
ster-

period

animals
response,

The
decreased
animals
to

lower

the
epinephrineto that
observed

lower

environmental

known.

The

animals
was
level
of liver

steroid.

DISCUssIoN.
adrenalectomized
normal

saline,

were

the
poor

until

The

8 hr ; with

salt

them

of
saline,
the
more
in body
temperature.

on

of

by

maintaining

of saline.

almost
a

conserve

on

longer

and
close

ture
vived

stores

normal
was

in normal
animals.
with
aldosterone

day

produced

test
brought
level of the ad-

effect.

the

glycogen

to levels
near
induced
glycosuria

sive

was strengthened
treatment
with

With

liver

were

demonstrated

after

deprived
the
decline

maintained

same

saline

the
steroid-treated
due to a lower
initial

was

was

cold

instead

were

survived

with

This

to

water

cold

prednisolone

or

maintained

response
presumably

treatment,

the

tap

Rats

norepi-

Treatment

catecholamine
back
to the

on

rats

animals
rapid

three

levels

although

cortisone

shortly
before
the
the glucose
response
while

glucose

obtained.

of

renalectomized

of all
to

energy.

exposing

the
norepiineffective.

response

complicated,

picture

single

FFA

for

more

of
was

administration

restored

nephrine.
The

metabolic

; acute

dexamethasone

the
lipolytic
action
while
aldosterone

contrast,

cold-exposure

115

II

MECHANISMS.

However,

animals

of approximately

heat

these

almost
if

untreated,
pretreatment
had

under
the

200

as fast
animals

cal/hr,

less

adrenalectomized
with
a single
virtually

conditions

no

as untreated
were

protec-

; such

ani-

rats.
given

daily

doses

of

steroids,

fective

as

might

aldosterone

the

abling
the
tam
their

other

that

in

dosterone

is

sodium,

thus
in

corticoids

has

at.

who

(1960),
vigor

tomized

and

sodium

and

levels

the

fall

summary,
that the
rats

cholamines.

to

and
FFA
for additional

mobilize

unable

to

temperature

cose

ability

pioerect,
and

FFA.

is
in

of
shiver

the

poor

to

cate-

3
and
to cold,

4.
are

such

as

substrates
needs
fuel.

body

heat

until

animals
the

blood

strengthened

tables
exposed

rapidly
the

to

meet
the
metabolic

conserve
of

the

and

not

unreasonable
to posof cold on adrenalec-

energy
to

drops

Pretreatment

volume

concept

glucose
bodies

the
did

end-organs

presented
animals,

unable

increased

chloride

attributed

the

This

by the
data
Adrenalectomized

strict,

be

of

plasma

deoxycor-

that

in plasma

may

adrenalecof

contrast,
and

et
main-

level

doses

Swingle
in

low

In

it is not
lethal
effect

responsiveness

stores

activity

in

time.

prednisolone

sodium

action

mineralo-

by

that

normal

plasma

tomized

also

and

acetate
of

counteract
pressure.
In
tulate

longer

reported

al-

renal

chloride.

ticosterone

its

gluco-

despite

are
while

of

found

efen-

glucocorticoids

requiring
been

as
in

to maincold.
One

manner,

through

actions

dogs

4)

animals
in the

acute

effective

just

(table

the

some

difference

tains

was

steroids

adrenalectomized
body
temperature

speculate

effective
on

ET

MAICKEL

116

with
animals

and

to

of their
They
are
and

death

their
occurs.

steroids
to

re-

vasocon-

mobilize

glu-

Vol.

AL.

157

REFERENCES
Cussou.,
N. V., LONOLBY,
R. W. AND Ros, J. H.:
The
determination
of glycogen
in liver
and
muscle
by the use of anthrone
reagents.
J. Biol.
Chem.
220:
583-593,
1956.
Doz.s,
V. P. : A relation
between
non-esterified
fatty
acids
in plasma
and
the metabolism
of
glucose.
J. Clin. Invest.
35: 150-154,
1956.
GUILLEMIN,
R., CLAYTON,
G. W., Lipscosss,
H. S.
AND SMITH,
J. D. : Fluorometric
measurement
of
rat plasma
and adrenal
corticosterone
concentration.
J. Lab. Clin. Med. 53: 830-832,
1959.
MAICKEL,
R. P.
BRODIE,
B. B. : Interaction
of
drugs
with
the pituitary-adrenocortical
system
in the production
of the fatty
liver.
Ann. N.Y.
Acad. Sci. 104:
1059-1064,
1963.
MAICK.EL,
R. P., MATUSSEK,
N., STERN,
D. N. AND
BRODIE,
B. B. : The sympathetic
nervous
system
as a homeostatic
mechanism.
I. Absolute
need
for sympathetic
nervous
function
in body
ternperature
maintenance
of cold-exposed
rats.
J.
Pharmacol.
Exp.
Therap.
157:
103-110,
1967.
RAMEY,
E. R. 4tm Gousrnzw,
M. S. : The adrenal
cortex
and
the
sympathetic
nervous
system.
Physiol.
Rev. 37: 155-195,
1957.
SAWYER,
M. E. M. AND SCHLOSSBERO,
T. : Studies
of homeostaais
in normal,
sympathectomized
and
ergotaminized
animals.
Amer.
J. Physiol.
104:
172-189,
1933.
Szcxzn,
J. : A note on the suprarenal
cortex
and
the transmission
of the activity
of the sympathetic
nerves
of the cat. J. Physiol.
(London)
95: 282-285, 1939.
SHAFRIR,
E. AND STEINBERG,
D. : The essential
role
of the adrenal
cortex
in the response
of free
fatty
acids,
cholesterol
and
phospholipids
to
epinephrine
injection.
J. Clin.
Invest.
39: 310319, 1960.
SHAFRIR,
E., SUSSMAN,
K. E. AND STEINBERG,
D.:
Role
of the pituitary
and the adrenal
in the
mobilization
of free fatty acids and lipoproteins.
J. Lipid Res. 1 : 459-465,
1960.
SWINGLE,
W. W., DAVANZO,
J. P., GLENISTER
D.,
WAGLE,
G., Osnomn,
M. AND ROWEN,
R. : Eects
of mineraloand
glucocorticoids
on
fasted
adrenalectomized
dogs subjected
to electroshock.
Proc. Soc. Exp. Biol. Med.
104:
184-188,
1960.