Effects of Fasting During Pregnancy on Maternal and

Fetal Weight and Body Composition in

Well-Nourished and Undernourished Rats1'2
SALLY ANN LEDERMAN ANDPEDRO ROSSO
Institute of Human Nutrition and Department of
Pediatrics, College of Physicians and Surgeons,
Columbia University, New York, NY 10032

A 50% food restriction during preg

nancy has been shown to reduce fetal size
in the rat (1). In spite of the adverse ef
fect on the fetus, changes in maternal net
body weight and body composition are
the same in pregnant and nonpregnant
rats during food restriction, indicating
that the fetus cannot parasitize the tissues
of an undernourished mother to main
tain normal growth.
On the other hand when fasted for 2
days late in pregnancy, previously wellfed rats have been reported to lose more
weight than do fasted nonpregnant rats
and fetal growth is maintained (2).
The apparent difference between food
restriction and fasting may be explained
in several ways. For example, adaptation
to fasting during pregnancy may be sig
nificantly different from adaptations to
food restriction so that fasting results in
greater mobilization of maternal nutrient
stores. Also, the greater weight losses ob
served in fasted pregnant rats may be
1823

caused by changes in body composition,

such as a greater loss of water, and may
not signify greater nutrient mobilization
for fetal use in the pregnant animals.
Finally, well-fed animals fasted for 2 days
in late pregnancy may be well above their
prepregnancy weight and may there
fore still contain body fat gained in early
pregnancy. This excess weight and fat
may be a prerequisite for increased ma
ternal nutrient mobilization during fast
ing and for the consequent maintenance
of fetal growth while the mother loses
weight. If so, undernourished rats would
show a different response to fasting dur
ing pregnancy than is shown by wellnourished rats.
The purpose of the present study was
Received for publication 20 February 1981.
'Supported in part by NIH grant KO 4 HD 00116 (Research
Career Development Award).
1The work reported in this paper was a portion of a Ph.D. disser
tation submitted to Columbia University by S. A. Lederman in 1960.
An abstract of part of this work has appeared in Fed. Proc.3S(3):871,
1979.

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ABSTRACT
The effect of a 2-day fast on fetal and maternal weight and com
position was determined in ad libitum-fed and food-restricted pregnant and nonpregnant rats. Fasting between days 17 and 19 of gestation resulted in a greater
loss of net maternal body weight in ad libitum-fed pregnant than in nonpregnant rats and also a greater loss of body fat. In contrast, food-restricted preg
nant rats, also fasted from day 17 to day 19 of gestation, maintained their net
body weight and body fat during the fast as did nonpregnant rats fasted for the
same length of time. Fetal weight was not significantly reduced by fasting in
the ad libitum-fed rats but was reduced by 25% in the previously food-re
stricted rats. The results demonstrate that prior maternal nutritional status
strongly influences the effects of fasting on the fetus and that maternal nutrient
stores are not mobilized for fetal utilization even when fetal growth is markedly
impaired.
J. Nutr. Ill: 1823-1832, 1981.
INDEXING KEY WORDS
fasting body composition fetal weight

1824

LEDERMAN

to determine the changes in fetal and ma

ternal weight and in maternal composi
tion during fasting in well-nourished and
undernourished
pregnant rats and to com
pare these to changes in similarly fed
groups of nonpregnant rats to determine
whether maternal nutrient stores are mo
bilized to aid fetal growth during fasting
and whether pre-existing undernutrition
alters the effects of fasting on maternal
weight and fetal growth.

Female Sprague-Dawley
rats (Holtzmann Co., Madison, WI) weighing 220260 g on day 0 (day on which preg
nant animals were found to be "sperm
positive") were received on day 4 of gesta
tion. The rats were individually caged on
arrival and maintained in a temperatureregulated room with a 12-hour light-dark
cycle. Water was available at all times.

SOT FASTED
AD LIBITUM
FASTED
PREGNANT
NOT FASTED
FOOD RESTRICTED
FASTED

NOT FASTED
AD LIBITUM
FASTED
NON PREGNANT
NOT FASTED
FOOD RESTRICTED
FASTED

17

EXPERIMENTAL

19

DAYS

Fig. 1 Timetable shows division of rats into ad

libitum-fed and food-restricted groups and then into
not fasted and fasted groups.

All animals were fed a standard labora

tory diet ad libitum until day 5 (RQ Rat
and Mouse Diet, Zeigler Bros. Inc., Gard
ners, PA). On day 5,10 pregnant and eight
nonpregnant rats were killed and the re
maining animals were divided into foodrestricted and ad libitum-fed subgroups as
shown in figure 1. On day 17, seven or
more rats from each diet group were
killed and the remaining animals in each
of the four groups were distributed be
tween not fasted and fasted subgroups.
The not-fasted groups continued on their
prior intake. The fasted groups were to
tally deprived of food. The remaining ani
mals (7-11 per group) were killed on
day 19.
Ad libitum-fed animals were given a
weighed amount of excess food and
the uneaten portion was collected and
weighed about every 2 days. Food-re
stricted rats were given their pre-weighed
daily food ration every morning. All foodrestricted
animals received
the same
amount of food, about 50% of the mean
daily intake of the ad libitum-fed preg
nant group. Food was weighed to the
nearest 0.1 g. Animals were weighed
to the nearest 0.1 g about twice weekly
and before being killed. Only animals
bearing 10 2 fetuses were included in
the pregnant groups.
Animals were killed with an overdose
of chloroform. The maternal .stomach and
intestines were excised and discarded
after they had been stripped of visible fat
which was replaced in the carcass. In
pregnant rats, the fetal sac was severed at
the base of the placenta and weighed. The
fetuses were then freed of the extra-fetal
tissues
which were discarded.
The
carcass, liver, pooled fetuses and pooled
placentas of each animal were weighed
separately.
All retained
tissues were
frozen.
Carcass analysis was performed on four
or five animals which were representa
tive of their group with respect to weight.
The tail was removed from the carcass,
weighed and discarded prior to carcass
analysis. Carcass composition is based on
the carcass weight without the tail.
On the day of analysis, the carcass was
cut with shears into several pieces

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MATERIALS AND METHODS

AND ROSSO

1825

FASTING DURING PREGNANCY IN THE RAT

and then ground (with fur and bone)

through the coarse and fine blades of a
meat grinder until it was the consistency
of finely ground meat. Triplicate
0.8-2 g samples of the ground carcass
were analyzed for water by drying to con
stant weight at 95-100.
For the determination of carcass fat con
tent, duplicate samples of 1-3 g of ground
carcass were weighed into 125-ml Erlenmeyer flasks. The tissue was dispersed in
20 ml of vacuum-redistilled hexanes,
warmed for 20 minutes or more on a steam
bath and filtered through sintered glass.
The solvent was evaporated off and the fat
content was determined. Fetal water con
tent was determined by drying four or
more fetuses per dam to constant weight
at 95-100.
Statistical
methods.
The Students'
data weretanalyzed
by the
two-tailed

RESULTS

TABLE 1
Food intake and body weight in ad libitum-fed

and food-restricted

rats

Total body wt
Day 5

Pregnantad
libNonpregnantad

Day 12

Day 17

Day 19

Carcass wt

Day
killed

2245
4a258
43223223225
324
2a229
324
300

_256
266 2a(Fasted)205

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Food intake. Pregnant rats consumed

an average of 25 g of food per day during
the ad libitum feeding period. Intake of
nonpregnant rats averaged 19 g/day.
During the restriction period, food-re
stricted rats consumed 12 g per day or 47%
of the intake of the ad libitum-fed preg
nant group. This represented 62% of the
intake of ad libitum-fed nonpregnant rats.
Body weight. Changes in total body
weight and carcass weight are shown in
table 1. The change in net maternal
weight from day 5 to day 19 in fasted and
not fasted rats is shown in table 2. Net
maternal weight is the total weight minus
the weight of fetuses and placentas. Since
different animals provided the data for
net weight changes to day 17 and to day
19, the changes are not exactly additive.
Ad libitum-fed nonpregnant and preg
test (3). Results were considered sig
nant
rats gained net weight between days
nificantly different if the P value was 0.05
5 and 17, although pregnant rats gained
or less.

2222
261
2227
libPregnantrestrictedNonpregnantrestricted2472462482532412412392402482522502402402392'5223353
253
230
2207
251
266
2(Fasted)204
2*>204

253
132243277289287256258252252256258236240237
2"189
256
241
2"(Fasted)197

223
226

2"200
228
4"183
224
203
2"(Fasted)101081088677101186851719195171919171919171919

533231332242351
43223242209
3318

1Mean SEM. " Significantly different from corresponding nonpregnant group (P < 0.05). Only
carcass weights were compared.
b Significantly different from corresponding ad libitum fed group
(P < 0.05). Only carcass weights were compared.

1826

LEDERMAN

AND ROSSO

TABLE 2
Weight changes in ad libitum-fed
wt
change before
fasting, days
17'+58.2
5-

Pregnant
libNonpregnant
ad

2.2(7)-l

libPregnant
ad

(8)-11.1

wt change,
17-19Notdays

2.0"

15.5 2.5"

fastedg+9.0

1.3"

1.0"(11)+4.8

(10)-23.3
0.9(7)-14.4
*

(6)+

2.5"

19'Not

maternal wt change,
days 5-

fasted+67.2

.
0.7"

4.6"

15.7 1.6"

(15)-23.3

(6)+26.8

0.9-(7)-24.5

4.5"(6)-11.7

3.2"(7)-35.6

1.0(15)-21.1

&2(7)-

3.1
(11)-36.6

(10)-9.8

(6)-0.6

1.3(8)-2.2
13.4

0.6(11)-21.1

1.2'(6)Fasted-23.9

0.5-(8)Net

0.9(H)-2.2

0.5-(8)Net

1.2(6)Fasted-42.5

14.62.5(6)Fasted+

1.4(8)

restrictedMaternal
1Weight of fetuses and placentas subtracted in pregnant groups.
* Weight change during fasting in pregnant groups = (net weight
change days 5 to 19) - (net weight change days 5 to 17). n for pregnant groups therefore equals the degrees of freedom. Weight change in
nonpregnant groups = total weight change during fasting.
3 Mean SEM; numbers in parentheses indicate the number of animals.

Significantly different from corresponding pregnant group (P < 0.05).

b Significantly different from corresponding food-restricted
group (P < 0.05).

substantially
more. If they were not
fasted, both nonpregnant and pregnant
ad libitum-fed rats gained additional net
weight between days 17 and 19. During
2 days of fasting, the nonpregnant group
had a moderate weight loss and ended the
fast below their day 5 weight.
During fasting, ad libitum-fed pregnant
rats also lost net weight. After fasting how
ever, their net body weight remained
above their day 5 weight. Although the
pregnant rats lost more net weight when
fasted than the nonpregnant rats, they still
ended the fast heavier than the nonpreg
nant rats did.
The 50% food-restricted
nonpregnant
rats lost weight during the period of food
restriction. When fasted 2 days, they lost
additional weight and they ended the fast
well below their day 5 weight.
Food-restricted
pregnant rats lost net
weight at a rate similar to restricted nonpregnant rats. When fasted between day
17 and 19, previously
food-restricted
pregnant rats also lost additional weight
and, like the nonpregnant restricted rats,
ended the fast well below their day 5
weight. Thus, restricted pregnant and
nonpregnant rats were the same weight at
the end of the fast.
The results indicate that during a 2-day
fast, previously ad libitum-fed pregnant

rats lose more net maternal weight than

similar nonpregnant rats, as previously re
ported (2). Previously restricted, fasted
pregnant and nonpregnant rats have the
same overall net weight loss.
Body composition.
Maternal carcass
composition was determined on days 5,17
and 19 in fasted and not fasted groups,
to establish whether fasting affected body
composition differently in pregnant or
nonpregnant rats and whether prior food
restriction influenced body composition
changes during fasting. The body compo
sition data are shown in tables 3 and 4.
Fasting did not significantly change
percent composition in previously wellfed nonpregnant and pregnant rats (table
3). However, the absolute amount of
water, fat and lean dry tissue tended to be
lower after fasting in previously well-fed
nonpregnant
rats and, in the pregnant
group maternal carcass water and fat de
creased significantly.
Percent lean dry tissue increased and
percent fat decreased significantly during
fasting in previously restricted pregnant
rats (table 4). In restricted nonpregnant
rats only the decrease in percent carcass
fat was statistically significant.
In restricted nonpregnant rats, carcass
water, fat and lean dry tissue were lower
after fasting but only the decrease in

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1.8"(8)Gross

2.5-

rats fasted for 2 days or not fasted

maternal wt change,
17-191JNot
days

fasted+26.8

(8)+4.8

1.7(6)-16.8

restrictedNonpregnant

or food-restricted

1827

FASTING DURING PREGNANCY IN THE RAT

Carcass composition

TABLE 3
changes during fasting in groups previously ad libitum fed1
Day 19

Day 5

65.2 0.8
10.9 1.3
23.9 0.7

Water, g
Fat, g
Lean dry, g

133.1 1.6
22.4 2.9
48.7 1.1

Nonpregnant
libWater,
%Fat,
%Lean
%Water,
dry,

Not fasted

Fasted

64.812.522.7162.431.456.80.70.60.2"1.5b1.7e1.3C
64.910.624.5144.823.854.60.50.90.62.6abc2.50.6C
65.112.022.9155.528.654.80.50.70.3"1.6*2.0"0.4C

-10.7
-4.8
-0.2

ad

gFat,
gLean
dry, g63.8

1.613.0

0.923.2

0.9128.6

4.026.1

1.746.7

1.663.9

1.010.5

0.525.6

0.5137.2

3.622.6

0.854.9

1.1e63.610.725.7139.523.41.21.30.1e3.03.155.8
0.6e63.910.026.1129.120.11.30.7e0.83.01.4e52.6
1.6e-8.1-2.5-2.3

1n = 4, all groups; results expressed as means SEM.

(P < 0.05).
" Significantly
different from corresponding
c Significantly different from day 5 value (P < 0.05).

carcass fat was statistically significant. In

restricted pregnant rats, fasting caused a
significant decrease in carcass water and
fat but lean dry tissue was not signifi
cantly changed. Absolute changes in
carcass components in the fasted animals
between day 17 and 19 derived from the
mean values for the carcass composition
in the group killed before fasting, on day
17, and the group killed after fasting, on
day 19, are shown in tables 3 and 4. The
values indicate that well-fed pregnant rats
lost more body fat when fasted than
nonpregnant rats did (table 3).
In previously food-restricted rats (table
4), what appeared to be a slightly greater
weight loss during fasting in pregnant rats
compared to nonpregnant rats was due to
a greater loss of carcass water in the preg
nant rats. Previously restricted pregnant
rats lost 10.5 g carcass water during fast
ing whereas nonpregnant rats lost only 4
g. Pregnant and nonpregnant rats lost
similar amounts of fat and lean dry tissue
stores, however (table 4).
During fasting, total body water was
reduced in both of the pregnant groups

a Significantly different from day 17 value

nonpregnant
group value (P < 0.05).

but in the restricted pregnant rats, total

body water fell below that of the ad libitum-fed nonpregnant fasted groups and
was not significantly different from re
stricted, fasted nonpregnant rats.
Liver weight. Liver weight decreased
in all fasted groups as shown in table 5.
The liver weight of both pregnant groups
remained above that of the correspond
ing nonpregnant group during fasting.
Well-fed pregnant and nonpregnant rats
had a comparable loss of liver weight dur
ing fasting, 3.8 and 3.2 g respectively.
In previously restricted rats, pregnant
rats lost 1.4 g of liver weight during fast
ing and nonpregnant rats lost 2.2 g
(table 5).
Fetal and placental weight. Table 6
shows the fetal and placental weights of
previously ad libitum-fed or food-re
stricted fasted and not fasted rats.
On day 19, fasted rats previously fed ad
libitum had fetuses weighing 92% and
placentas weighing 86% of the values for
not fasted rats. The differences were not
significant by the two-tailed Student's ttest. In previously well-fed rats, placental

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Pregnant ad lib
Water, %
Fat, %
Lean dry, %

Day 17

Change
during
fasting

1828

LEDERMAN

AND ROSSO

TABLE 4
Carcass composition changes during fasting in groups previously food restricted1
Day 19
Day 17

Not fasted

Change
during
fasting

Fasted

Pregnant restricted
Water, %
Fat, %
Lean dry, %

Nonpregnant
restrictecWater,
%Fat,
%Lean
%Water,
dry,

67.6 0.6"
7.1 1.1"
25.3 0.5"

69.6 0.8cd
2.4 0.8aed
28.0 0.4acd

134.2 2.4M
14.2 2.6d
51.3 0.2"

133.9 2.0M
14.0 2.2"
50.2 1.2

123.7 2.0a1
4.4 1.6acd
49.8 1.5"

1.010.4

1.09.0

1.125.9

0.8e125.9

1.226.3

0.8e122.6

4.0"20.2

2.451.2

1.165.1

gFat,
gLean

dry, g163.3

-10.5
-9.8
-1.5

1.9"17.4

2.2C50.2

2.566.65.328.1118.69.550.11.11.1"0.42.0"20acd1.1-4.0-10.7-1.1

1n = 4, all groups; results expressed as means SEM. a Significantly different from day 17 value
(P < 0.05).
b Significantly different from corresponding nonpregnant group value (P < 0.05).
c Sig
nificantly different from day 5 value (P < 0.05).
d Significantly different from corresponding
ad
libitum group (P < 0.05).

weight increased significantly during the

fasting period and was not significantly
lower than in not fasted rats.
Food restriction to day 19 reduced fetal
and placental weight to 88 and 78% of con
trol, respectively, but the decrease in fetal
weight was not statistically significant.
Fetal weight has been shown to be re
duced significantly by food restriction
continued to day 21 (1). To determine if

the slowed rate of fetal growth established

during food restriction could be main
tained during a fast, the fetal weights on
day 19 of previously food-restricted fasted
and not fasted rats were compared. Food
restricted fasted rats had fetuses only 74%
of controls, significantly lighter than the
fetuses of the not fasted restricted rats.
Restricted rats also had significantly
smaller placentas than ad libitum-fed

TABLE 5
Liver weight in ad libitum-fed

and food-restricted

pregnant and nonpregnant

19Pregnant

Day
511.8

ad lib
Nonpregnant ad lib
Pregnant restricted
Nonpregnant restrictedDay

rats'

1716.2

0.4
12.7 0.6Day 11.9
11.0
10.0

fastedg16.9

0.6""
0.2ab
0.2"^
8.7 0.1e"
0.4
13.0 0.2
0.4e
11.2 0.2^
9.6 O.!"0"6
0.2aeNot 9.3 0.2aeFasted12.4
7.7 O.lcdeLiver

wt change
duringfasting-3.8

-3.2
-1.4
-2.2

1Results expressed as means SEM. " Significantly different from corresponding day 5 value
(P < 0.05).
b Significantly different from corresponding nonpregnant group value (P < 0.05).
c Sig
nificantly different from not fasted value (P < 0.05).
d Significantly different from day 17 value
(P < 0.05).
e Significantly different from corresponding ad libitum group value (P < 0.05).

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Water, g
Fat, g
Lean dry, g

67.2 1.1M
7.1 1.3d
25.7 0.4d

1829

FASTING DURING PREGNANCY IN THE RAT

TABLE 6
Fetal and placental weight and fetal composition in ad libitum-fed

and food-restricted

rats1

Day 19
Day 17

Not fasted

Fetal wt, g
Ad libitum fed
restrictedPlacental
Food

0.04
0.78
0.080.56

0.15"
2.25
0.070.77

wt, g
Ad libitum fed
restrictedFetal
Food

0.02
0.03"90.5
0.45

0.04a
0.02ab89.0
0.60

90.3

0.2
0.42.57

88.6

0.2
0.6a2.36

0.12
0.05ab0.66
1.91
0.03a
0.02""89.1
0.55

88.8

0.1"
0.2"

1Results expressed as means SEM. a Significantly different from day 17 value (P < 0.05).
nificantly different from corresponding ad libitum group value (P < 0.05).

rats both at day 17 and day 19. Similarly,

the placentas of fasted restricted rats were
significantly smaller than those of fasted,
previously well-fed rats.
Placental weight at day 19 was lower in
animals restricted from day 5 of gestation
than it was in animals fasted from day 17
to 19. Although 2-day fasted rats had
somewhat smaller placentas at day 19
than did unfasted rats, the differences
were not significant in either the previ
ously well-fed or food-restricted groups.
Fetal body composition. The effect of
fasting on fetal body water is shown in
Table 6. Fetal percent body water de
clined between days 17 and 19 in fasted
and not fasted rats whether previously
well fed or food restricted. In spite of the
large effect on fetal weight, restriction fol
lowed by fasting did not significantly alter
fetal percent body water at day 19.
DISCUSSION
This study confirms previous observa
tions that in well-fed rats a normal rate of
fetal growth can be maintained during a
2-day fast. During this fast pregnant ani
mals lose more weight (conceptus sub
tracted) than similarly fed nonpregnant
animals. Carcass constituent losses are
slightly higher in pregnant than in nonpregnant previously well-fed fasted rats
and include greater fat and water losses
with less lean tissue loss.

b Sig

This finding is consistent with the wellestablished observation that fasted preg
nant rats (4) and humans (5, 6) mobilize
fat more readily than when not pregnant.
It is believed that this more rapid re
sponse to starvation is designed to en
able fetal use of glucose. Well-fed preg
nant rats lost less lean dry tissue during
fasting than did nonpregnant
rats, sug
gesting that total glucose utilization may
have been less in the pregnant animals.
The larger amount of food in the alimen
tary tract of pregnant rats or their liver glycogen stores may also have supplied the
glucose needed by the conceptus during
the fast. However, liver weight decreased
similarly during fasting in pregnant and
nonpregnant rats.
Although previously well-fed pregnant
rats lost more total weight (conceptus sub
tracted) than nonpregnant rats during fast
ing, changes in carcass composition and
carcass weights were not markedly dif
ferent. These findings indicate that body
components not included in the carcass,
i.e., the liver and intestines, were re
sponsible for the greater weight loss in
the pregnant group. The liver data show
that there was no large difference in liver
weight losses in the two ad libitum-fed
groups when fasted. Therefore,
the
greater net weight loss of the pregnant
group was due to a larger loss in intestinal
weight, probably due to the fact that
pregnant rats had more intestinal contents

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% Water
Ad libitum fed
Food restricted0.81

Fasted

1830

LEDERMAN

shown to induce natriuresis and water

loss during fasting in humans (10). Preg
nant rats experience
a much greater
fall in blood glucose during fasting than
do nonpregnant rats (11) and they would
be expected to have a more marked gluca
gon response.
It is especially noteworthy that the loss
of carcass fat in previously restricted preg
nant and nonpregnant rats during fasting
was more than twice the amount lost by
the previously well-fed fasted animals.
This surprising finding requires explana
tion.
Restricted rats began fasting with their
adaptive mechanisms already taxed. Dur
ing fasting, they were exposed to a lifethreatening
stress and biological
re
sponses were probably pushed to their
limits. After 3 days of fasting, pregnant
rats have higher urinary epinephrine
and norepinephrine
excretion than nonpregnant rats (12). On the other hand,
blood glucose is much lower during fast
ing in rats that are pregnant (2). Stressstimulated
lipolytic hormone
release
(cortisol, placental lactogen, growth hor
mone, glucagon) is probably also en
hanced when food restriction precedes
fasting (13-16). Strangely, although more
fat was mobilized, it did not substitute
for other energy sources because total
carcass nutrient losses were greater in
both groups of restricted fasted rats com
pared to ad libitum-fed rats when fasted.
This result may also bea consequence of
heightened glucagon secretion since ex
ogenous glucagon has been shown to in
crease protein catabolism in fasted rats
(14). It is also known that energy expend
ing activity is increased in food restricted
rats during a fast (17). In addition, absorp
tion of nutrients from the intestines would
have ceased earlier in fasting in previ
ously food-restricted
rats, necessitating
greater use of body stores.
Although fetal weight in fasted re
stricted rats was reduced significantly to
only 85% of the not fasted food restricted
value, placental weight was not signifi
cantly reduced by fasting. Fasted re
stricted rats had placentas 91.6% of the
weight of the placentas of unfasted re
stricted rats. On the other hand, in not

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at the beginning of the fasting period and

therefore would defecate a larger amount.
The different alimentary losses make it
appear that pregnant rats lose more tissue
weight than nonpregnant
rats during
fasting, but they do not. Clearly, caution
must be used to interpret weight changes
when net weight without intestinal con
tents is not determined.
Since ad libitum-fed
pregnant
rats
gained more weight than nonpregnant
rats prior to the fast, they ended the fast
with a higher carcass weight than nonpregnant fasted rats in spite of their
greater weight loss during fasting.
In previously food-restricted rats, there
is a greater weight loss during fasting in
pregnant rats than in nonpregnant
rats
but carcass analysis shows that the dif
ference is due to greater decreases in
body water in the pregnant rats. Com
pared to nonpregnant rats, no additional
fat or lean dry tissue is lost during fast
ing in previously food restricted preg
nant rats.
The decrease in body water in the
fasted restricted
pregnant group may
have significance for fetal growth. It has
been shown that either food restriction or
a low protein diet reduces the blood vol
ume increase that normally occurs in
pregnancy (7). Uterine blood flow is also
reduced (8). Our total body water meas
ures at day 17 (table 3) indicate that ad
libitum-fed pregnant rats had higher body
water content than ad libitum-fed nonpregnant rats. On day 17, restricted preg
nant rats (table 4) did not have a higher
body water than well-fed nonpregnant
rats (table 3).
If blood volume follows total body
water, placental
perfusion
and fetal
growth could be severely compromised
in the fasted restricted pregnant rats
where total body water was markedly re
duced. For example, in humans it has
been found that mothers of small babies
have a smaller plasma volume and total
body water in late pregnancy
than
mothers of large babies (9).
The reduction
in total body water
which occurred during fasting in both
pregnant groups may be mediated by
glucagon secretion. Glucagon has been

AND ROSSO

FASTING DURING PREGNANCY IN THE RAT

nancy might be always ill-advised, it

would be particularly undesirable in
mothers previously not well nourished.
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fasted groups placental weight at day 17

and day 19 was significantly lower in re
stricted rats than in ad libitum-fed rats,
but fetal weight was not. These results
show that fetal and placental growth al
though related, are not totally interde
pendent. As other workers have demon
strated, the tissue growing most rapidly is
most affected by a period of undernutrition (18). During the period when fasting
was imposed in the present study, fetal
growth was much more rapid than pla
cental growth and therefore it was more
affected by the fast.
This study demonstrates that fasting
like food restriction significantly reduces
fetal growth in undernourished rats. The
accelerated metabolic adaptations to
starvation which characterize pregnancy
do not protect the fetuses of undernour
ished mothers. It is likely that these adap
tations are effective in previously wellfed animals because maternal stores of
glucogenic materials are sufficient to pro
vide for fetal needs temporarily if the
mother can metabolize fat. These adapta
tions do not always protect the fetus.
Fasting reduces fetal growth in food-re
stricted animals, in rats fasted more than
2 days (19, 20), or in rats fasted so early in
pregnancy that maternal metabolism has
not yet made the adaptations typical of
late pregnancy, and maternal protein, fat
and glycogen stores have not yet in
creased. In each of these cases the lack of
appropriate maternal substrates, par
ticularly fat and carbohydrate, may render
useless the maternal adaptive processes
which would otherwise protect the fetus
from periods of short-term maternal fasts.
These considerations are of major signifi
cance
pregnancy,
where (21)
the
belief for
that human
"accelerated
starvation"
may prevent fetal growth retardation is
easily misapplied. Nevertheless, due to
the biological differences between rat and
human pregnancy, the implications of the
findings for human gestation cannot be
stated with certainty.
In a broader context these results in
fasted rats show that prior nutritional
status may alter the effect of a specific nu
tritional treatment on the mother or on the
fetus. Thus, although fasting during preg

1831

1832

LEDERMAN

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vation-induced
activity in dietary obese,
dietary lean and normal weight rats. Behav.
Biol. 24,220-228.

AND ROSSO
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normal
genase
Nature

M. M. (1972) Effects of starvation on

development of b-OH butyrate dehydroactivity in foetal and newborn rat brain.
New Biol. 236, 140-141.

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Assan, R. & Marliss, E. B. (1977) Fetal meta
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(1978) Is there a risk of "accelerated starva
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27, 463.